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PROCEEDINGS 

of the 

Biological Society of 
Washington 



VOLUME 82 



WASHINGTON 
PRINTED FOR THE SOCIETY 






y 



/ 



A^ 



r^sSTHSOTv^ 



JUN 1 1970 



EDITOR 
Roger F. Cressey 



All correspondence should be addressed to 
the Biological Society of Washington, Smith- 
sonian Institution, Washington, D.C. 20560 



Allen Press Inc. 
Lawrence, Kansas 66044 



OFFICERS AND COUNCIL 

OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 

FOR 1969-1970 

OFFICERS 

President 
JOSEPH ROSEWATER 

Vice President 
MEREDITH L. JONES 

Secretary 
RICHARD C. RANKS 

Treasurer 
HARALD A. REHDER 



COUNCIL 

Elected Members 



THOMAS E. BOWMAN 
K. C. EMERSON 



JAMES A. PETERS 
GEORGE A. LLANO 



CLAYTON E. RAY 



ShV. . 
INSllil 



{^Ay2? W 



TABLE OF CONTENTS 

Volume 82 

Becklund, Willard W. The role of the national parasite collec- 
tion in veterinary parasitology 603-610 

Black, Joe B. A new crawfish of the genus Hobbseus from 

Mississippi (Decapoda, Astacidae) 193-200 

Campden-Main, Simon M. The identity of OUgodon cyclurus 
(Cantor, 1839) and revalidation of OUgodon brevicauda 
( Steindachner, 1867) (Serpentes: Colubridae) 763-766 

Choate, Jerry R. Taxonomic status of the shrew, Notiosorex 
(Xenosorex) phillipsii Schaldach, 1966 (Mammalia: In- 
sectivora) 469^76 

Cowan, Richard. The national collections as biological standards 
579-584 

Crabill, R. E., Jr. A new floridan Cryptops with key to the 
state's species (Chilopoda: Scolopendromorpha: Cryptopi- 
dae) 201-204 

Cressey, Roger F. Five new parasitic copepods from California 

inshore fish 409-428 

Cressey, Roger F. Hatschekia pacifica new species (Copepoda: 
Caligoida) a parasite of the sand bass, Faralebrax nebulifer 
( Giard ) ^ 843-846 

Cutler, Edward B. New species of Sipuncula from the Western 

North Atlantic 209-218 

Dawson, C. E. Citharichthys abbotti, a new flatfish (Bothidae) 

from the southwestern Gulf of Mexico 355-372 

Dawson, C. E. Paragunnellichthys fehlmanm, a new gobloid 

fish ( Microdesmidae ) from the Indian Ocean 373-380 

Deevey, Georgiana B. Pelagic ostracods (Myodocopa halo- 

cyprididae) from the North Atlantic off Barbados 799-824 

Elbel, Robert E. Chapinia elbeli Tendeiro, a synonym of 

Chapinia fasciati Elbel (Mallophaga: Menoponidae) 489-490 

Fleming, Laurence E. Use of male external genitalic details as 
taxonomic characters in some species of Palaemonetes 
( Decapoda, Palaemonidae ) 443-452 

Foster, Nancy M. New species of spionids (Polychaeta) from 
the Gulf of Mexico and Caribbean Sea with a partial revision 
of the genus Priorwspio 381-400 

Gates, G. E. Contributions to a revision of the earthworm 

family Lumbricidae V. Eisenia zebra Michaelsen, 1902 453-460 

Gloyd, Howard K. Two additional subspecies of North Amer- 
ican crotalid snakes, genus Agkistrodon 219-232 

Gray, W. Scott, Jr., and John C. McCain. The taxonomic status 
of Mandibulophoxus gilesi Barnard, 1957 (Crustacea: Am- 
phipoda) 189^192 

iv 



Halpern, Jerald A. Biological investigations of the deep sea. 46. 

The genus Litonotaster ( Echinodermata, Asteroidea) 129-152 

Halpern, Jerald A. Biological investigations of the deep sea. 50. 
The validity and generic position of Pentagonaster parvus 
Perrier (Echinodermata, Asteroidea) 503-506 

Higgins, Robert P. Indian Ocean Kinorhyncha 2. Neocentro- 

phyidae, a new homalorhgid family 113-128 

Hobbs, H. H., III. A new genus and two new species of 

entocytherid ostracods from Alabama and Mississippi 167-170 

Hobbs, Horton H., Jr. Two new species of the crayfish genus 
Procambarus (Decapoda, Astacidae) with keys to the mem- 
bers of the Spiculifer Group 329-348 

Hobbs, Horton H., Jr., and Edward T. Hall, Jr. New crayfishes 

from Georgia (Decapoda Astacidae) 281-294 

Hobbs, Horton, H., Jr., and Joseph F. Fitzpatrick, Jr. A new 
crayfish of the genus FaUicambarus from Tennessee (De- 
capoda, Astacidae ) 829-836 

Hobbs, Horton H., Jr., and Margaret Walton. New entocytherid 

ostracods from Tennessee and Virginia 851-864 

Hoffman, Richard L. Studies on spiroboloid millipeds. VII. A 
remarkable new genus and subfamily of the Spirobolelhdae 
from Vera Cruz, Mexico 177-188 

Hope, W. D., and D. G. Murphy. Phaptothyreus typicus n. g., 
n. sp., an abyssal marine nematode representing a new family 
of uncertain taxonomic position 81-92 

Hope, W. D., and D. G. Murphy. Syringonomus typicus new 
genus, new species (Enoplida: Leptosomatidae ) a marine 
nematode inhabiting arenaceous tubes 511-518 

Hotton, Nicholas III. Vertebrate fossil collections — ^A frag- 
mentary document 579-584 

Kenk, Roman. Freshwater triclads ( Turbellaria ) of North 

America. I. The genus Planaria 539-558 

Klappenbach, Miguel A. A new species of Eupera (MoUusca, 

Pelecypoda) from Haiti 835-828 

Kornicker, Louis S. Bathyconchoecia deeveyae, a highly orna- 
mented new species of Ostracoda (Halocyprididae) from the 
Peru-Chile Trench System 403-408 

Lewis, Jackson E. Reversal of asymmetry of chelae in Calappa 

Weber, 1795 (Decapoda: Oxystomata) 63-80 

Manning, Raymond B. Notes on the Gonodactylus section of the 
family Gonodactylidae (Crustacea, Stomatopoda ) , with de- 
scriptions of four new genera and a new species 143-166 

Manning, Raymond B. The postlarvae and juvenile stages of 
two species of Pseudosquillopsis (Crustacea, Stomatopoda) 
from the Eastern Pacific region 525-538 

Manning, Raymond B. Automation in museum collections 671-686 



Manning, Raymond B. Notes on the west American nephro- 

pidean lobster, Nephropsis occidentalis Faxon 865-870 

McCain, John C, A new species of caprellid (Crustacea: 

Amphipoda) from Oregon 507-510 

McCain, John C. Familial taxa within the Caprellidae (Crus- 
tacea: Amphipoda ) 837-842 

Miller, Milton A., and Welton L. Lee. A new isopod, Idotea 
(Pentidotea) kirachanskii, from Central California (Crus- 
tacea) 789^798 

Minckley, W. L., and Glady L. Lytle. Notropis xanthicara, 
a new cyprinid fish from the Cuatro Cienegas Basin, North- 
Cential Mexico 491-502 

Nichols, Frederic H. Tenoma kitsapensis, a new genus and 
species of the family Polynoidae (Polychaeta) from Puget 
Sound (Washington) 205^208 

Parkes, Kenneth C. The Red-throated Ant-tanager {Habis 

fuscicauda) in Panama and Colombia 233-242 

Peters, James A. A note on the generic names Cyclagras Cope 

and Lejosophis Jan (Reptilia: Serpentes) 847-850 

Pettibone, Marian H. Review of some species referred to 

Scalisetosus Mcintosh (Polychaeta, Polynoidae) 1-30 

Pettibone, Marian H. Remarks on the North Pacific Harmothoe 
tenehricosa Moore (Polychaeta, Polynoidae) and its associa- 
tion with asteroids ( Echinodermata, Asteroidea) 31-42 

Pettibone, Marian H. The genera Polyeunoa Mcintosh, Hol- 
olepidella Willey, and three new genera ( Polychaeta, Polynoi- 
dae) 43-62 

Pettibone, Marian H. The genera Sthenelanella Moore and 

Euleanira Horst (Polychaeta, Sigalionidae ) 429-438 

Pettibone, Marian H. Australaugeneria pottsi, new name for 
Polynoe longicirrus Potts, from the Maldive Islands (Poly- 
chaeta: Polynoidae ) 519-524 

Price, Charles E., and Eric S. McClellan. The monogenean 
parasites of African fishes. IX. A new genus, Gussevstrema, 
recovered from the gills of Terapon jarbua (Forskal) from 
South Africa 171-176 

Price, C. E., E. S. McClellan, A. Druckenmiller, and L. G. 
Jacobs. The monogenean parasites of African fishes, X. Two 
additional Dactylogyrus species from South African Barbus 
hosts 461-468 

Ritterbush, P. C. Art and science as influences on the early 

development of natural history collections 561-578 

Roper, Clyde F. E., C. C. Lu, and Katharina Mangold. A new 
species of lllex from the western Atlantic and distributional 
aspects of other lllex species (Cephalopoda: Oegopsida) 295-322 



Rosewater, Joseph. Malacological collections — development and 

management 663-670 

Sakai, Tune. Two new genera and twenty-two new species 

of crabs from Japan 243-280 

Schmitt, Waldo L. Colombian freshwater crab notes 93-112 

Schwartz, Albert. A new species of large Diploghssus (Sauria: 

Anguidae) from Hispaniola 777-788 

Setzer, Henry W. A new jackal (Canis adustus) from the 

Sudan 401-402 

Shetler, Stanwyn G. The herbarium: past, present, and future .. 687-758 

Shipp, Robert L., and Ralph W. Yerger. A new puffer fish, 
Sphoeroides parvus, from the western Gulf of Mexico, with a 
key to species of Sphoeroides from the Atlantic and Gulf 
coasts of the United States 477-488 

Smith- Vaniz, William F. A new species of Meiacanthus ( Pisces : 
Blenniidae: Nemophidinae ) from the Red Sea, with a review 
of the Indian Ocean species 349-354 

Springer, Victor G., and Robert E. Trist. A new cHnid fish 

species, Paraclinus fehlmanni, from Ecuador 323-328 

Stone, Benjamin C. Fandanus decus-montium, a new species 

from the Solomon Islands 439-442 

Sturtevant, William C. Does anthropology need museums? 619^650 

Wetmore, Alexander. Descriptions of additional forms of birds 

from Panama and Colombia 767-776 

Yochelson, Ellis L. Fossils — The how and why of collecting 

and storing 585-602 

Zusi, Richard L. The role of museum collections in ornithologi- 
cal research 651-662 



vii 



BIOLOGICAL SOCIETY OF WASHINGTON 
PROCEEDINGS 

1041st Meeting — 17 February 1969 

NINETIETH ANNUAL MEETING 

President Joseph Rosewater called the 1041st meeting and 90th 
Annual Meeting of the Society to order at 2:08 p.m. in Room 43, MNH. 
Fourteen members were present. 

Recording Secretary Stanwyn Shetler read the minutes of the 89th 
Annual Meeting of 27 June 1968, and they were approved as read. 

Corresponding Secretary Richard Banks reported that during the year 
20 new members had been added, while three former members, whose 
names he read, had died {see list attached to Council Minutes for 17 
Feb. 1969). 

Treasurer Harald Rehder gave his financial report, and Henry Roberts 
reported that the Auditing committee had found the Treasurer's books 
in order as of 16 February. The Treasurer's report was accepted. 

Corresponding Secretary Banks reported the results of the recent 
election of officers, noting that 73 ballots had been returned, of which 
4 had not been marked. 

Roger Cressey, Editor of the Proceedings, reported that a record 
number of papers (62) and pages (760) had been published during 
1968 and that 1969 promises to be another big year. He said that the 
papers given at the October 1968 symposium would go to the printer 
in April and be published in August. 

President Rosewater, on behalf of the Society, expressed thanks to Dan 
Cohen for his fine work in organizing the fall symposium. He asked 
the members present to think about topics for another symposium, be- 
cause various members thought another should be held this year. At 
his request, Shetler explained briefly his idea that a symposium on local 
flora and fauna might be appropriate, in view of the great attention paid 
to local flora and fauna in the early days of the Society. 

President Rosewater proposed that the Society accept the constitutional 
amendments as expressed in the letter that went to all members, and a 
formal motion to that effect was made by K. C. Emerson and seconded 
by Rehder. The motion passed unanimously. Rosewater reminded the as- 
sembled members that the adoption of these amendments has stream- 
lined the Society's organization by abolishing the office of Second Vice 
President and merging the offices of Recording and Corresponding 
Secretaries. The new Secretary is Richard Banks, and Rosewater thanked 
Shetler for his services to the Society as Recording Secretary. 

The meeting adjourned about 2:30 p.m. 

Stanwyn G. Shetler 
Recording Secretary 



IS 



QH 

1 82, pp. 1-30 29 May, 1969 

B4X 

^^H PROCEEDINGS 

OF THE 
BIOLOGICAL SOCIETY OF WASHINGTON 

REVIEW OF SOME SPECIES REFEMIED TO 

SCALISETOSUS MCINTOSH 

(POLYCHAETA, POLYNOIDAE) 

By Marian H. Pettibone 
Smithsonian Institution, Washington, D. C. 

In connection with an extended review of the polynoid gen- 
era, based on a study of the type-species, it was found that 
Scalisetosus Mcintosh ( 1885) has been used for a heterogenous 
group of species. The genus has served to include species with 
setae as transparent as crystal and the neurosetae character- 
ized by the presence of a basal semilunar cusp or pocket, al- 
though this particular feature was not shown on the figure of 
the neurosetae of the type-species, S. ceramensis, by Mcintosh 
(1885, pi. lOA, fig. 14). Any species equipped with this pe- 
culiar type of neurosetae has been placed in Scalisetosus, re- 
gardless of other characters. Saint- Joseph (1899) proposed the 
new genus Adyte for three species ( Polynoe pellucida Ehlers, 
Hermadion assimile Mcintosh, and H. echini Giard) having 
the peculiar type of neurosetae, separating them from S. cera- 
mensis, which lacks the basal semilunar cusps. Mcintosh 
(1900) was responsible for changing the diagnosis of Scali- 
setosus to include the species referred to Adyte by Saint- 
Joseph. Subsequently, Adyte was abandoned and was synony- 
mized with Scalisetosus by Fauvel (1914, p. 47). 

During a visit to the British Museum of Natural History in 
May 1967, I was able to examine the unique type of Scalise- 
tosus ceramensis and to verify that the neurosetae indeed do 
lack the basal semilunar cusps and that the species therefore 
lacks one of the key characters that has been attributed to the 
genus. The purpose of this paper is to report on the re-exami- 
nation of the holotype of S. ceramensis, as well as some of the 



1— Prog. Biol. Soc. Wash., Vol. 82, 1969 



JUN2& 1969 



2 Proceedings of the Biological Society of Washington 



other species of polynoids that have been referred to Scalise- 
tosus. Additional species were available for study in the British 
Museum of Natural History (BMNH), the Zoological Museum 
Amsterdam (ZMA), Swedish State Museum Natural History 
(SSMNH), the Zoological Museum Hamburg (ZMH), the 
Zoological Museum Berlin (ZMB), and the United States Na- 
tional Museum ( USNM ) . An effort was made to separate the 
species in more meaningful ways by emending Adyte Saint- 
Joseph and by establishing four new genera. The study is 
summarized by providing a key to the genera previously in- 
cluded under Scalisetosus. 

The following genera and species are included in the study: 



Scalisetosus Mcintosh 

Adyte Saint-Joseph, EMENDED 

Subadyte new genus 



S. ceramensis Mcintosh 
A. assimilis (Mcintosh) 

S. pellucida (Ehlers), new 
combination 

S. papillifera (Horst), new 
combination 

S. mjoehergi (Augener), 
new combination 

P. crinoidicola (Potts), 
new combination 

P. tentaculata (Horst), 
new combination 

P. praelongus 

(Marenzeller), new 
combination 

A. rutilans (Grube), new 

combination 
A. michaelseni new species 

This study was aided in part by a grant from the National 
Science Foundation (NSF GB-1269). I am indebted to R. W. 
Sims and D. George for their assistance during my visit to the 
British Museum (Natural History) and for the loan of speci- 
mens. Thanks are also extended to S. van der Spoel of the 
Zoological Museum Amsterdam, to R. Olerod of the Swedish 



Paradyte new genus 



Pottsiscalisetosus new genus 



Austrahugeneria new genus 



Review of Scalisetosus 3 

State Museum of Natural History, to G. Hartwich of the Zoo- 
logical Museum Berlin, and to G. Hartmann-Schroder of the 
Zoological Museum Hamburg for the loan of specimens. The 
manuscript benefited from the suggestions of F. A. Chace, Jr. 
and M. L. Jones, both of the Smithsonian Institution. 

FAMILY POLYNOIDAE MALMGREN 
Genus Scalisetosus McIntosh, 1885 

Type-species: S. ceramensis Mcintosh, 1885, by monotypy. Gender: 
masculine. 

Diagnosis: Body flattened, elongate-oval, tapered posteriorly, com- 
posed of about 40 segments. Elytra 16 pairs, arranged on segments 2, 4, 
5, 7, alternate segments to 23, 26, 29, 32, and 35. Prostomium bilobed, 
with distinct cephalic peaks, with 2 palps and 3 antennae; ceratophore 
of median antenna inserted in anterior notch; ceratophores of lateral an- 
tennae distinct, inserted ventrally. Tentacular segment (I) with 2 pairs 
tentacular cirri, with few setae, without facial tubercle. Buccal segment 
( II ) without dorsal nuchal fold; with long ventral buccal cirri. Parapodia 
subbiramous. Notopodia with long projecting acicular lobes. Notosetae 
much stouter than neurosetae, clear, without spinous rows but with few 
stout spines. Neuropodia with long projecting presetal acicular lobes, 
from which tips of acicula project, and shorter rounded postsetal lobes. 
Neurosetae slender, with faint spinous regions, with tips slightly fal- 
cate, entire ( lower ones ) and bifid ( upper ones ) ; without basal semilunar 
pockets or cusps. Dorsal cirri with short, cylindrical cirrophores. Dorsal 
tubercles nodular. Ventral cirri short, tapered. Nephridial papillae dis- 
tinct, short, cylindrical. 

Scalisetosus ceramensis Mcintosh 
Fig. 1 

Scalisetosus ceramensis Mcintosh, 1885, p. 103, pi. 13, fig. 7, pi. lOA, 

figs. 13, 14. 
^Scalisetosus ceramensis. — Horst, 1917, p. 98. 

Material examined: Challenger station 195A south of Ceram Island, 
4° 31' S, 129° 57' E, 659 meters, volcanic mud, 29 September 1874. — 
Holotype (BMNH 85: 12: 1: 85) Sihoga station 267, east of Great Kei 
Island, 5° 54' S, 132° 56.7' E, 984 meters.— 1 specimen (ZMA 1294). 

Description: As in generic diagnosis. Neither specimen is complete, 
although the holotype is tapered posteriorly and appears to be nearly 
complete, consisting of 40 segments. There are two dorsal transverse 
ciliated bands per segment. All of the elytra and the styles of the dorsal 
cirri are missing. Most of the notosetae are provided with 4 spines each 
but the number varies from 2 to 5. The nephridial papillae begin on 
segment 9. The pharynx was not extended and was not examined. 

Distribution: East Indies. In 659 to 984 meters. 



SWiinisuNif t MAY ZH 1969 



/: 



4 Proceedings of the Biological Society of Washington 




Figure 1. Scalisetosus ceramensis (Holotype, BMNH 85: 12: 1: 85): 
a, Anterior end, dorsal view; elytra, palps, styles of antenna, tentacular 
and dorsal cirri missing; eyes faded; b, same, ventral view; ventral buccal 
cirri of segment II missing; c, elytrigerous parapodium, anterior view, 
elytron missing; d, cirrigerous parapodium, posterior view, style of dorsal 
cirrus missing; e, notosetae; f, upper and lower neurosetae. 



Review of Scalisetosus 5 

Genus Adyte Saint-Joseph, 1899, Emended 

Type-species: Hermadion assimile Mcintosh, 1874, herein designated. 
Gender: feminine. 

Remarks: Adyte Saint- Joseph was proposed for three species: Polynoe 
pellucida Ehlers (designated as type-species of Subadyte new genus; see 
below), Harmadion assimile Mcintosh [= Adyte assimilis (Mcintosh) 
Saint- Joseph] and H. echini Giard [referred to Scalisetosus assimilis ( Mc- 
intosh) by Fauvel, 1923]. 

Diagnosis: Body flattened, elongate, subrectangular, tapered posterior- 
ly, composed of numerous segments (up to 74). Elytra 15 pairs, arranged 
on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, and 32, leaving 
large number of posterior segments without elytra (up to 42). Elytra 
delicate, smooth, without tubercles or fringes of papillae but with scat- 
tered sensory papillae. Prostomium bilobed, with lobes rounded anteriorly 
or with slight indication of cephalic peaks, with 2 palps and 3 an- 
tennae; ceratophore of median antenna inserted in anterior notch, cera- 
tophores of lateral antennae distinct, inserted ventrally. Tentacular seg- 
ment (I) with 2 pairs tentacular cirri, without setae; without facial 
tubercle. Buccal segment (II) without dorsal nuchal fold; with long 
ventral buccal cirri. Parapodia subbiramous. Notopodia with short coni- 
cal acicular lobes. Notosetae as stout as or stouter than neurosetae, nearly 
smooth, with scattered, closely appressed spinous rows along convex 
border. Neuropodia with diagonally truncate, rounded presetal lobes 
containing acicula (acicula not extending to tips of lobes) and short, 
rounded postsetal lobes. Neurosetae all similar (lower ones shorter), 
with basal semilunar cusps or pockets and faint distal spinous rows, with 
tips hooked, minutely bifid. Dorsal cirri with cylindrical cirrophores and 
with styles long, smooth, somewhat inflated below terminal filament. 
Dorsal tubercles inconspicuous. Ventral cirri short, subulate, Nephrid- 
ial areas bulbous but without definite papillae. 

Adyte assimilis (Mcintosh) 
Figs. 2, 3 

Hermadion assimile Mcintosh, 1874, p. 194; 1876, p. 387, pi. 70, figs, 4-6. 
Adyte assimilis. — Saint-Joseph, 1899, p. 167, pi. 6, figs, 1-3. 
Scalisetosus assimilis. — Mcintosh, 1900, p. 377, pi. 30, fig. 15, pi. 33, 

fig. 6, pi. 40, figs. 20-22.— Fauvel, 1923, p. 74, fig. 27, g-i.— St0p- 

Bowitz, 1948, p. 10, fig. 6. 

Material examined: Porcupine Expedition, off Spanish coast, 110-293 
meters, on Echinus esculentus, 1870. — Syntype (BMNH 1921: 5: 1: 
499). St. Andrews, Scotland.— Syntype (BMNH 1921: 5: 1: 498). Clew 
Bay, Ireland, 9-11 meters, on Echinus. — 1 specimen (BMNH 1914: 12: 
1258). North Sea, 49° 35' N, 6° 47' W, 128 meters, 17 September 1895, 
R. Meyer, collector.— 1 specimen (SSMNH 1262). 

Description: As in generic diagnosis. The specimen from Clew Bay 
is complete, consisting of 74 segments, 30 mm in length, 3.5 mm in width. 



6 Proceedings of the Biological Society of Washington 




Figure 2. Adyte assimilis (SSMNH 1262): a, Anterior end, dorsal 
view; elytra removed; b, prostomium, dorsal view; c, cirrigerous para- 
podium, posterior view; d, elytrigerous parapodium, anterior view, ely- 
tron removed; e, notosetae; f, upper, middle and lower neurosetae. 



Review of Scalisetosus 




Figure 3. Adyte assimilis (BMNH 1914: 12: 1258): a, Elytrigerous 
parapodium from anterior region, anterior view; b, cirrigerous parapo- 
dium from anterior region, posterior view; c, notoseta from same; d, upper 
and lower neurosetae from same; e, cirrigerous parapodium from posteri- 
or region, posterior view; f, notosetae from same; g, upper and lower 
neurosetae from same. 



^ 



8 Proceedings of the Biological Society of Washington 

including parapodia, and 4.5 mm in width, including setae. The North 
Sea specimen has a continuous middorsal longitudinal dark band. There 
are two transverse dorsal ciliated bands per segment. The pharynx has 11 
pairs of soft papillae, the two lateral pairs being much smaller. 

Distribution: North Atlantic, North Sea, EngHsh Channel, Mediter- 
ranean. Dredged. Commensal with echinoderms (echinoids). 

Subadyte new genus 

Type-species: Polynoe pellucida Ehlers, 1864. Gender: feminine. 

Diagnosis: Body flattened, elongate-oval, tapered posteriorly, com- 
posed of about 40 segments. Elytra 15 (or 16) pairs, arranged on seg- 
ments, 2, 4, 5, 7, alternate segments to 23, 26, 29, 32 (and sometimes 
34). Elytra delicate, transparent, with large vesicular soft papillae. Pro- 
stomium bilobed, with lobes rounded anteriorly, with or without distinct 
cephahc peaks, with 2 palps and 3 antennae; ceratophore of median an- 
tenna inserted in anterior notch; ceratophores of lateral antennae dis- 
tinct, inserted ventrally. Tentacular segment (I) with 2 pairs tentacular 
cirri, without setae; without facial tubercle. Buccal segment (II) with 
small nuchal fold; with long ventral buccal cirri. Parapodia subbiramous. 
Setae crystal clear. Notopodia with short projecting acicular lobes. Noto- 
setae subequal in width to neurosetae, with spinous pouches and blunt 
tips which may be slightly notched. Neuropodia with long projecting 
presetal acicular lobes, from which the tips of the acicula project, and 
shorter rounded postsetal lobes; with or without distal cirriform append- 
age. Neurosetae all similar (except lower ones shorter), with basal semi- 
lunar pockets and distal spinous regions, with tips hooked, bifid. Dorsal 
cirri with cylindrical cirrophores, bulbous basally; styles long, tapering, 
papillate. Dorsal tubercles inconspicuous. Ventral cirri short, subulate. 
Nephridial papillae distinct, small. 

Subadyte pellucida (Ehlers), new combination 
Fig. 4 

Polynoe pellucida Ehlers, 1864, p. 105, pi. 2, fig. 10, pi. 3, figs. 5, 7-13, 

pi. 4, figs. 1-3. 
Hermadion fragile Claparede, 1868, p. 383, pi. 5, fig. 2; 1870, p. 380, 

pl. 2, fig. 2. 
Adyte pellucida. — Saint-Joseph, 1899, p. 170, pl. 6, figs. 4-5. 
Scalisetosus communis. — Mcintosh, 1900, p. 372, pl. 26, fig. 6, pl. 27, 

fig. 12, pl. 30, fig. 9, pl. 33, fig. 7, pl. 34, fig. 1, pl. 40, figs. 17-19. 

Not Lysidice communis Delle Chiaje, 1841 (fide Claparede, 1870, p. 

380). 
Hermadion pellucidum. — Alaejos y Sanz, 1905, p. 31, pl. 4, fig. 7, pl. 5, 

figs. 1-7. 
Scalisetosus pellucidus. — Fauvel, 1914, p. 47; 1923, p. 74, fig. 27, a-f. — 

Monro, 1930, p. 48. 
Scalisetosus fragUis.— Day, 1962, p. 631; 1967, p. 59, fig. 1.7, g-k. 



Review of Scalisetosus 




C 

Figure 4. Subadyte pellucida (ZMA 1287): a, Anterior end, dorsal 
view, right palp regenerating; all tentacular cirri missing except for lower 
right one; elytra missing; b, cirrigerous parapodium, posterior view; c, 
elytrigerous parapodium, anterior view; d, notosetae; e, upper and lower 
neurosetae. 



10 Proceedings of the Biological Society of Washington 

Material examined: Tafelbaai, near Cape Town, South Africa, 5-8 
meters, M. Weber, collector, 23 July 1894.— 30 specimens (ZMA 1287) 
Isle of Man, west of Peel, 27-55 meters.— 1 specimen (BMNH 1923: 5 
501). Bressay Sound, Scotland, 18 meters. — 2 specimens (BMNH 1921 
5: 1: 505). Naples, H. A. Bayliss, collector. — 4 specimens (BMNH 1919 
11: 6: 30). Discovery station MS 82, Saldanha Bay, South Africa, 7-14 
meters, 6 September 1926.— 1 specimen (BMNH 1930: 10: 254). South 
Africa, J. H. Day, collector.— 1 specimen (BMNH 1961: 9: 837). 

Description: As in generic diagnosis. The body is extremely fragile. 
An almost complete specimen from Tafelbaai has 40 segments, 15 pairs 
of elytra, a length of 22 mm, a width of 6 mm, including parapodia, and 
10 mm, including setae. The prostomium lacks distinct cephalic peaks. 
The styles of the antennae and dorsal cirri have short claviform papillae. 
The elytra are oval, transparent, finely dotted with small claviform 
sensory papillae and some larger, oval soft papillae. The notosetae are 
as stout as or sHghtly stouter than the neurosetae; they are curved, with 
denticled pouches along the convex side ( 6-20 ) ; the tips are blunt, with 
a small slit. The projecting neuropodial presetal acicular lobe has the tip 
of the aciculum projecting (without a fiHform appendage, as in S. papil- 
lifera; see below). There are two transverse dorsal ciliated bands per 
segment. The small nephridial papillae begin on segment 6. The pharynx 
has 11 pairs of soft papillae, the two lateral pairs being small. 

Distribution: Adriatic, Mediterranean, North Atlantic, North Sea, 
English Channel, Bay of Bengal, South Africa. Dredged. Found among 
algae, commensal with echinoderms (asteroids, ophiuroids). 

Remarks: Mcintosh (1900) used Scalisetosus communis (Delle Chiaje) 
for this species but Lysidice communis Delle Chiaje is considered inde- 
terminable. Day (1962, p. 631; 1967, p. 59) replaced Scalisetosus pel- 
lucidum (Ehlers, 1864) with S. fragilis (Claparede, 1868), since he con- 
sidered the former name preoccupied. However, Polynoe pellucida Ehlers 
(1864) precedes Polynoe pellucida (Dyster, in litt.) Johnston (1865, p. 
117). Thus, the latter is the junior homonym, not the former. 

Subadyte papillifera (Horst), new combination 
Fig. 5 

Scalisetosus papuliferous Horst, 1915, p. 17; 1917, p. 99, pi. 21, figs 
2-4.— Monro, 1928, p. 469. 

Material examined: Sihoga station 43, Anchorage off Pulu Sarasso, 
Postilion Islands, coral, 4-5 April 1890.— Syntype (ZMA 1286). Tahiti.— 
5 specimens (BMNH 1928: 1: 11: 30). Marianas, Lagoon west of Saipan, 
May to June, 1949, P. E. Cloud, collector.— 6 specimens (USNM 26055 
to 26058). Caroline Islands, Ifahk Atoll, August 1953, D. P. Abbott, col- 
lector.— 1 specimen (USNM 22977). 

Description: As in generic diagnosis. The body is extremely fragile. 
The syntype has 38 segments and 15 pairs of elytriphores. Some speci- 
mens show a characteristic mottled pattern dorsally. The elytra are deli- 



Review of Scalisetosus 



11 




Figure 5. Subadyte papillifera (USNM 26057): a, Anterior end, 
dorsal view; styles of median antenna, left and upper right tentacular 
cirri missing; elytra missing; b, cirrigerous parapodium, posterior view; 
c, elytrigerous parapodium, anterior view; d, distal tip of neuropodium 
of same; e, notosetae; f, upper, middle and lower neurosetae. 



12 Proceedings of the Biological Society of Washington 

cate, transparent, with some scattered (9-20), thick, elongate-oval soft 
papillae (pi. 21, figs. 3, 4, in Horst, 1917). The prostomial lobes are sub- 
triangular anteriorly, with indistinct or with sHghtly developed cephalic 
peaks. The setae are crystal clear. The notosetae are slightly more 
slender than the neurosetae; they are short to long, with spinous pockets 
along the curved edge; the tips are blunt and may be shghtly notched. 
The projecting presetal acicular lobes of the neuropodia have the tips of 
the acicula projecting and a cirriform appendage below the aciculum. 
There are two transverse dorsal ciliated bands per segment. The small 
nephridial papillae begin on segment 6. The pharynx has 11 pairs of 
papillae, the two lateral pairs being small. 

Distribution: Central Pacific, Malay Archipelago. Associated with 
corals. Littoral. 

Subadyte mjoebergi ( Augener ) , new combination 
Fig. 6 

Scalisetosus mjohergi Augener, 1922a, p. 12, fig. 2. 

Material examined: West Austraha, 45 miles W.S.W. Cape Jaubert, 22 
meters, E. Mjoberg, collector, 1 June 1911.— Holotype (SSMNH 594). 

Description: As in generic diagnosis. The holotype has 41 segments 
and 16 pairs of elytriphores (last pair on segment 34); it is 12 mm in 
length, 3 mm in width, including parapodia, and 5 mm in width, includ- 
ing setae. The elytra are delicate, transparent, with numerous (up to 
30), large oval soft papillae and small clavate sensory tubercles on distal 
parts of large papillae, as well as on surface of elytra (figs. 2, 2a, b, in 
Augener, 1922a). The prostomial lobe has distinct cephalic peaks. The 
setae are crystal clear. The notosetae are subequal in width to the 
neurosetae; they are short to long, with spinous pockets along the curved 
edge; the tips are blunt and sHghtly notched. The projecting presetal 
acicular lobes of the neuropodia have the tips of the acicula projecting 
(may be broken) and a cirriform appendage. There are two transverse 
dorsal cihated bands per segment. The small nephridial papillae begin on 
segment 6. The pharynx has 11 pair of papillae, the two lateral pairs 
being small. 

Distribution: Southwest Australia. In 22 meters. 

Remarks: Scalisetosus australiensis Benham (1915), also from Aus- 
traha, should perhaps be referred to Subadyte; the incomplete descrip- 
tion was based on a damaged anterior fragment of 28 segments. 

Paradyte new genus 

Type-species: Polynoe crinoidicola Potts, 1910. Gender: feminine. 

Diagnosis: Body flattened, elongate-oval, tapered posteriorly, seg- 
ments about 40. Elytra 15 pairs, arranged on segments 2, 4, 5, 7, alter- 
nate segments to 23, 26, 29, and 32. Elytra deHcate, smooth, without 
tubercles or fringes of papillae (may have scattered sensory papillae). 
Prostomium bilobed, with lobes rounded, subtriangular anteriorly, with or 



Review of Scalisetosus 13 

without distinct cephalic peaks, with 2 palps and 3 antennae; ceratophore 
of median antenna inserted in anterior notch; ceratophores of lateral 
antennae distinct, inserted ventrally. Tentacular segment (I) with 2 
pairs tentacular cirri, without setae; without facial tubercle. Buccal 
segment (II) without dorsal nuchal fold; with long ventral buccal cirri. 
Parapodia subbiramous. Notopodia with short conical acicular lobes. 
Notosetae stouter than neurosetae, curved, sabre-like, with few spines 
along convex border, with tips notched or entire. Neuropodia with diag- 
onally truncate, subtriangular presetal lobes containing aciculum (acicu- 
lum may or may not extend to end of lobe) and short rounded postsetal 
lobes. Neurosetae of two kinds: supra-acicular slender, with basal semi- 
lunar pockets and more distal elongate spinous regions and slightly 
hooked, bifid tips; subacicular much stouter, with basal semilunar pock- 
ets and short falcate smooth tips. Dorsal cirri with cylindrical cirrophores, 
bulbous basally; styles long, smooth, tapering. Dorsal tubercles incon- 
spicuous. Ventral cirri short, subulate. Nephridial papillae distinct, short, 
bulbous. 

Paradyte crinoidicola ( Potts ) , new combination 

Fig. 7 

Polynoe crinoidicola Potts, 1910, p. 337, pi 18, fig. 10, pi. 20, fig. 30, 

pi. 21, figs. 39-41. 
Scalisetosus (Polynoe) crinoidicola. — Horst, 1917, p. 98, pi. 16, figs. 6-8, 

pi. 21, fig. 1. 
Scalisetosus crinoidicola. — Okuda, 1936, p. 564, fig. 3. 
Scalisetosus longicirrus. — Fauvel, 1953, p. 50, fig. 22, a-c. — Day, 1962. 

p. 631; 1967, p. 58, fig. 1.7, a-f.— [?] Not Polynoe (L.) longicirra 

Schmarda, 1861. 

Material examined: Siboga station 164, 1° 42.5' N, 130° 47.5' E, 
32 meters, sand, small stones and shells, 20 August 1899. — 1 specimen 
(ZMA 1285). Siboga station 167, 2° 35.5' S, 131° 26.2' E, 95 meters, 
22 August 1899.— 1 specimen (ZMA 1994). Inhaca Island, Delagoa 
Bay, South Africa, 25° 50' S, 32° 50' E, commensal on crinoid, Tropio- 
metra carinata. — 1 specimen (BMNH 1936: 1: 10). Caroline Islands, 
IfaHk Atoll, F. M. Bayer, D. P. Abbott, collectors, 1953.— 4 specimens 
(USNM 22972, 22973, 22976). 

Description: As in generic diagnosis. The body is extremely fragile 
and is dorsally darkly pigmented with hghter spots, with prominent 
transverse dorsal cihated bands, 2 per segment. The elytra are also darkly 
pigmented with hghter spots and have numerous sensory papillae. The 
prostomial lobes are subtriangular, without distinct cephalic peaks. The 
setae are bronze-colored. The notosetae are stouter than the neurosetae, 
curved, sabre-hke, with 1-5 widely spaced spines along the curved 
border, with tips notched. In the diagonally truncate presetal lobes of 
the neuropodia, the acicula do not extend to the tips of the lobes. The 
supra-acicular group of neurosetae are slender, slightly curved, with long 
spinous regions and bifid tips. The subacicular group of neurosetae are 



14 Proceedings of the Biological Society of Washington 




Figure 6. Subadyte mjoebergi ( Holotype, SSMNH 594 ) : a, Anterior 
end, dorsal view; styles of median antenna, upper tentacular and dorsal 
cirri missing; elytra missing; base only of extended pharynx shown; b, 
cirrigerous parapodium, posterior view; c, elytrigerous parapodium, an- 



Review of Scalisetosus 



15 




Figure 7. Paradyte crinoidicola (USNM 22972): a. Anterior end, 
dorsal view; styles of median and left lateral antennae and tentacular 
cirri missing; elytra missing; b, elytrigerous parapodium, anterior view, 
elytron missing; c, cirrigerous parapodium, posterior view; d, notosetae; 
e, supra-acicular neuroseta; f, upper, middle and lower subacicular neuro- 
setae; g, neurosetae from segment II. 



terior view; elytron missing; d, tip of neuropodium from same; e, noto- 
setae; f, upper, middle and lower neurosetae. 



16 Proceedings of the Biological Society of Washington 

much stouter, with tips entire and falcate. The pharynx was extended in 
the Siboga specimen from Station 164; it has 9 (dorsal) and 11 (ven- 
tral) papillae, the lateral pair being small. The small nephridial paillae 
begin on segment 8. 

Distribution: Maldives, Malay Archipelago, Caroline Islands, Japan, 
South Africa (Inhaca Island). Associated with echinoderms (crinoids). 

Remarks: Polynoe crinoidicola Potts (1910) was referred to Scalise- 
tosus longicirrus (Schmarda) by Fauvel (1953) and Day (1962, 1967). 
Polynoe (Lepidonotus) longicirrus Schmarda (1861) from Ceylon, how- 
ever, must be considered indeterminable. The latter species was stated 
to have 17 pairs of elytra. 

Paradyte tentaculata ( Horst ) , new combination 
Fig. 8 

Scalisetosus tentaculatus Horst, 1915, p. 18; 1917, p. 100, pi. 21, figs. 5-7. 

Material examined: Siboga station 274, east of Aru Islands, 5° 28.2' 
S, 134° 53.9' E, 57 meters.— Syntype (ZMA 1288). 

Description: As in generic diagnosis. The single complete specimen 
has 39 segments. There are 2 dorsal ciHated bands per segment. All the 
elytra are now missing (they were large, overlapping, and covered with 
small oval papillae, according to Horst). The prostomium has distinct 
cephahc peaks. The notosetae are stouter than the neurosetae, curved, 
with 1 to 3 spines along the curved border (mostly 2 spines); the tips 
are entire, not notched. In the projecting subtriangular presetal lobes of 
the neuropodia, the acicula extend to the tips. The supra-acicular group 
of neurosetae are slender, with long, faintly spinous regions and bifid 
tips. The subacicular group of neurosetae have short, faintly spinous re- 
gions and entire, sHghtly falcate tips. The pharjnix was not extended and 
was not examined. The small nephridial papillae begin on segment 8. 

Distribution: Malay Archipelago. In 57 to 69 meters. 

Remarks: As indicated by Horst (1915, 1917), P. tentaculata is closely 
allied to Paradyte levis ( Marenzeller, 1902, as Scalisetosus), new combi- 
nation, from Japan. In P. levis, some notosetae lack spines, others have 
single spines. Scalisetosus pacificus Izuka ( 1912 ) , also from Japan, ap- 
pears to be identical with P. levis. 

Pottsiscalisetosus new genus 

Type-species: Scalisetosus praelongus Marenzeller, 1902. Gender: 
mascuHne. 

Diagnosis: Body flattened, elongate-vermiform, tapered posteriorly, 
segments numerous (50-110 or more). Elytra numerous pairs (28 or 
more), arranged on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 
32, 33, 35, continuing on alternate segments to end of body (may be 
some irregularity after segment 39). Elytra soft, translucent, without 
tubercles or papillae. Prostomium bilobed, with lobes rounded, without 
distinct cephalic peaks, with 2 palps and 3 antennae; ceratophore of 



Review of Scalisetosus 



17 




SI 11 11 
d 



Figure 8. Paradyte tentaculata (Syntype, ZMA 1288): a, Anterior 
end, dorsal view; styles of median antenna, upper tentacular cirri and 
dorsal cirri missing; elytra missing; eyes faded; b, cirrigerous parapodium, 
posterior view; style of dorsal cirrus missing; c, elytrigerous parapodium, 
anterior view; elytron missing; d, notosetae; e, supra-acicular neuroseta; 
f, upper, and lower subacicular neurosetae. 



18 Proceedings of the Biological Society of Washington 




Figure 9. Pottsiscalisetosus praelongus (Holotype of Halosydna (?) 
willeyi Potts, BMNH 1924: 3: 1: 63): a, Anterior end, dorsal view; ely- 
tra missing; right lateral antenna regenerating; eyes faded (dotted); b, 
same, ventral view; c, two parapodia from middle region of body show- 
ing platelets on ventral bases of parapodia, ventral view; d, cirrigerous 



Review of Scalisetosus 19 

median antenna inserted in anterior notch; ceratophores of lateral anten- 
nae distinct, inserted terminally, on same level as median antenna. Ten- 
tacular segment (I) with 2 pairs tentacular cirri, without setae; without 
facial tubercle. Buccal segment (II) with middorsal bulbous area (no 
distinct flap); ventral buccal cirri only slightly longer than following 
ventral cirri. Parapodia subbiramous. Notopodia with short, subcorneal 
acicular lobes. Notosetae more slender than neurosetae, tapered to blunt 
tips, very finely serrated. Neuropodia long, distally deeply notched dor- 
sally and ventrally, forming subequal anterior and posterior rounded 
lobes. Neurosetae stouter than notosetae, all similar, slightly to distinctly 
hooked, finely serrated, with projecting semilunar cusps or pockets 
more basally. Dorsal cirri with long cylindrical cirrophores, with styles 
rather short, tapered. Dorsal tubercles inconspicuous. Ventral cirri short, 
subulate. Nephridial papillae inconspicuous, the nephridial area covered 
with platelets on ventral bases of parapodia. 

Pottsiscalisetosus praelongus (Marenzeller), new combination 
Fig. 9 

Scalisetosus praelongus Marenzeller, 1902, p. 575, pi. 3, fig. 11. — 

Imajima and Hartman, 1964, p. 39. 
Scalisetosus formosus Moore, 1903, p. 403, pi. 23, figs, 4-6. 
Halosydna zeylanica Willey, 1905, p. 250, pi. 1, figs. 12, 13. 
Halosydna (?) willeyi Potts, 1910, p. 340, pi. 21, figs. 44, 45. 

Material examined: Albatross station 3703, off Honshu, Japan, 57 
meters, 7 May 1900. — Holotype of Scalisetosus formosus Moore (USNM 
16165). J. S. Gardiner Sealark Expedition, Amirante, western Indian 
Ocean, 44-155 meters, 10 November 1905. — Holotype of Halosydna (?) 
willeyi Potts (BMNH 1924: 3: 1: 63). 

Description: As in generic diagnosis. The holotype of Halosydna (?) 
willeyi consists of 58 segments plus a regenerating posterior end of about 
8 segments, with a length of 21 mm, widths of 3 mm, including parapodia, 
and 4 mm, including setae. There are 2 transverse dorsal ciliated bands 
per segment. The prostomial antennae are short and stumpy. The tentac- 
ular and dorsal cirri are rather short. The neurosetae are all similar ex- 
cept the subacicular ones, which are slightly stouter and more distinctly 
hooked. The pharynx was not extended and was not examined. 

Distribution: Japan, Indian Ocean (Amirante, Ceylon). In 20 to 165 
meters. Commensal with echinoderms ( asteroids ) . 



parapodium from anterior region of body, posterior view; e, notoseta 
from same; f, upper, middle and lower neurosetae from same; g, ely- 
trigerous parapodium from middle region of body, posterior view; h, 
same, from posterior region of body; i, notosetae from same; j, upper, 
middle and lower neurosetae from same. 



20 Proceedings of the Biological Society of Washington 

Australaugeneria new genus 

Type-species: Polynoe rutilans Grube, 1878. Gender: feminine. 

Diagnosis: Body flattened, elongate-oval, tapered posteriorly, com- 
posed of about 40 segments. Elytra 15 pairs, arranged on segments 2, 4, 5, 
7, alternate segments to 23, 26, 29, 32. Elytra soft, translucent, without 
tubercles or papillae. Prostomium bilobed, with lobes rounded, with- 
out distinct cephalic peaks, with 2 palps and 3 antennae; ceratophore of 
median antenna inserted in anterior notch; ceratophores of lateral anten- 
nae distinct, inserted ventrally. Tentacular segment (I) with 2 pairs 
tentacular cirri, without setae; without facial tubercle. Buccal segment 
(II) without dorsal nuchal fold; notosetae lacking; neurosetate strongly 
hooked; ventral buccal cirri sMghtly longer than following ventral cirri. 
Parapodia subbiramous. Notopodia with projecting acicular lobes. Noto- 
setae stouter than or more slender than neurosetae, smooth or spinous. 
Neuropodia long, deeply notched dorsally and ventrally, forming subequal 
anterior and posterior rounded lobes; presetal neuropodial lobes of seg- 
ments II and III enlarged, hoodHke. Neurosetae of several kinds; stout 
hooks on segments II and III (and sometimes on IV); upper group slen- 
der, with wider basal regions and long, more slender spinous regions, with 
tips blunt; lower group stouter, with wider basal regions and short hooked 
tips (may be faintly spinous; may have small secondary tooth). Dorsal 
cirri with long cylindrical cirrophores, with styles long, filamentous. 
Dorsal tubercles inconspicuous. Ventral cirri short, subulate. Nephridial 
papillae short, wide, distinct in posterior half of body. 

Remarks: The type material of Scalisetosus hartmeyeri Augener, 
deposited in the Zoological Museum Hamburg, consisted of two lots: one 
lot (ZMA 7895) contained 29 syntypes (2 complete but with regenerat- 
ing posterior ends, 27 anterior fragments, 7 posterior ends and a few 
middle fragments); the other lot (ZMA 10087) contained a single speci- 
men (anterior and posterior fragments). Both lots were collected at the 
same station in Southwest Australia. On examination of this material, it 
was found to contain two species — a more slender one with spinous 
notosetae (25 anterior and 7 posterior ends from ZMA 7895) and a 
slightly stouter species with smooth notosetae ( anterior and posterior frag- 
ments from ZMA 10087 and 4 anterior fragments and a middle piece 
from ZMA 7895). The fact that one of the stouter specimens had been 
separated from the others suggests that Augener had noted some differ- 
ences. Both species are similar in most respects, the most remarkable 
character being the presence in segments II and III of modified neuro- 
setae in the form of golden hooks, enclosed in enlarged hoodlike presetal 
neuropodial lobes. A new genus, Australaugeneria, is herein proposed 
for the two species. Since Augener's description and figures agree more 
closely with the more numerous slenderer specimens, I have retained 
Scalisetosus hartmeyeri Augener for this group and from it selected a 
Lectotype and 24 Paralectotypes. The other group of 5 specimens is 
herein described as a new species, A. michaelseni. 



Review of Scalisetosus 



21 




FiGLiRE 10. Australaugeneria rutilans ( Paralectotype of S. hart- 
meyeri, ZMH 7895); a, Anterior end, dorsal view; styles of left lateral 
antenna and tentacular cirri missing; style of median antenna broken; 
b, elytrigerous parapodium from segment II, anterior view; c, neuropodial 
hooks from same; d, neuropodial hooks from segment III; e, elytrigerous 
parapodium from segment IV, anterior view; f, upper and lower neu- 
rosetae from same; g, notoseta from same; h, middle elytrigerous para- 
podium, anterior view; i, middle cirrigerous parapodium, posterior view; 
j, notoseta from same; k, upper and lower neurosetae from same. 



22 Proceedings of the Biological Society of Washington 

Augener ( 1913, p. 123 ) indicated that there were close similarities 
between his Scalisetosus hartmeyeri and Polynoe rutilans Grube (1878) 
from the Philippine Islands. The Holotype of the latter, deposited in the 
Zoological Museum Berlin (ZMB 1137) was examined. It consists of an- 
terior, middle and posterior fragments, totaling 37 segments ( 38, accord- 
ing to Grube). Although in poor condition, being hardened and brittle, 
it is sufficient to indicate general agreement between the holotype of 
Polynoe rutilans Grube and the lecto- and paralectotypes of Scalisetosus 
hartmeyeri Augener, particularly in regard to the spinous notosetae (not 
smooth, as in A. michaelseni) and the modified neuropodial hooks on 
segments II and III; this latter character was not mentioned by Grube. 

Australaugeneria rutilans (Grube), new combination 
Fig. 10 

Polynoe rutilans Grube, 1878, p. 37, pi. 2, fig. 5. 

Scalisetosus hartmeyeri Augener, 1913, p. 119, fig. 5, a-e, pi. 2, figs. 17, 
18 (part). 

Material examined: Lapinig, Philippine Islands, Professor Semper, col- 
lector, commensal with the alcyonarian Xenia. — Holotype (ZMB 1137). 
Hamburger Australian Exp., Sta. 14, Sharks Bay, Freycinet Reach, South- 
west Australia, 11 to 16 meters, 12 September 1905, W. Michaelsen and 
R. Hartmeyer, collectors. — Lectotype and 21 Paralectotypes of S. hart- 
meyeri (ZMH 7895); 3 Paralectotypes of S. hartmeyeri (USNM 38894). 

Description: As in generic diagnosis. The combined length of an an- 
terior and a posterior fragment, totaling 36 segments, is 7 mm, the width 
is 2 mm, including parapodia. The elytra are large, overlapping, covering 
the middorsum and parapodia. The prostomium has 4 small eyes on the 
posterior half. The neuropodial hooks occur on segments II and III; the 
neurosetae of setiger IV are transitional, being similar to those of the 
following segments but more strongly hooked. The notopodia form short 
conical lobes. The notosetae are relatively few in number (less than 10), 
curved, sabre-hke, with spinous rows (5-12) and bifid tips. The neuro- 
setae are relatively few in number (less than 10); the upper ones are 
slender, enlarged basally, with long spinous regions and blunt tips; the 
lower ones are stout, with short spinous regions and hooked tips; there 
may be a minute secondary tooth. The pharynx was not extended in any 
of the specimens. The short and wide nephridial papillae are distinct, 
beginning on segment 17. There are transverse ciHated bands, 2 per 
segment dorsally and one ventrally near the posterior border. 

Distribution: Philippine Islands, Southwest Austraha. In 11 to 16 
meters. Associated with alcyonarians. 

Australaugeneria michaelseni new species 
Figs. 11, 12 

Scalisetosus hartmeyeri Augener, 1913, p. 119 (part). 

Material examined: Hamburger Australian Exp., Sta. 14, Sharks Bay, 



Review of Scalisetosus 



23 




Figure 11. Australaugeneria michaelseni (Paratype, ZMH 10087): 
a, Anterior end, dorsal view; styles of antennae, tentacular and dorsal 
cirri missing; elytra missing; b, same ventral view, pharynx partially ex- 
tended; c, neuropodial hook from segment II; d, neuropodium from seg- 
ment III, anterior view; e, neuropodial hooks from same; f, neurosetae 
from segment IV; g, notoseta from same; h, elytrigerous parapodium 
from segment V. 



Freycinet Reach, Southwest Australia, 11 to 16 meters, 12 September 
1905, W. Michaelsen and R. Hartmeyer, collectors. — Holotype and 3 
Paratypes (ZMH 10087); Paratype (USNM 38893). 

Description: As in generic diagnosis. The holotype, consisting of an 
anterior fragment of 14 segments and a posterior fragment of 7 seg- 
ments (the latter all with cirrophores, corresponding perhaps to seg- 
ments 33 to 39), has a length of 6 mm and width of 3 mm, including 
parapodia. An incomplete paratype of 25 segments has a length of 11 mm 



24 Proceedings of the Biological Society of Washington 




Figure 12. Australaugeneria michaelseni ( Paratype, ZMH 10087 ) : a, 
Cirrigerous parapodium from middle region, posterior view; style of dor- 
sal cirrus missing; b, elytrigerous parapodium from middle region, ante- 
rior view; c, notosetae from same; d, upper and lower neurosetae from 
same. 



and a width of 4 mm, including parapodia. No elytra remain attached. 
The styles of the antennae, tentacular and dorsal cirri are all missing. 
No eyes are visible on the prostomium (perhaps faded). The neuro- 
podial hooks occur on segments II, III, and IV; on the latter segment, 
a transitional type of neuroseta occurs in addition to the hooks. The 
notopodia form inflated basal lobes with digitiform acicular lobes ex- 
tending to near the distal tips of the neuropodia. The notosetae are few 
in number (about 12), stout, spike-like, smooth, tapering to blunt, en- 
tire tips (may be worn). The neurosetae are few in number (less than 
10); the upper ones are more slender, with stouter basal portions and 
long spinous regions with blunt tips; the lower neurosetae are stout, 
with short spinous regions and slightly hooked tips. The pharynx was ex- 



Review of Scalisetosus 25 

tended on one of the paratypes; it is extended ventrally, at right angles 
to the body; the distal end is encircled with 9 pairs of papillae and has 
2 pairs of hght colored jaws. The short and wide nephridial papillae 
are distinct, beginning on segment 17. There are 2 prominent transverse 
dorsal ciliated bands per segment, continuing on the medial bases of the 
elytriphores and cirriphores; ventrally there are 1 to 2 bands along the 
posterior border of the segment and on the ventral bases of the parapodia 
medial to the ventral cirri. 

Distribution: Southwest Australia. In 11 to 16 meters. 

Remarks: The main differences between the two species of Austral- 
augeneria are indicated below: 

A. rutilans (Grube) A. michaelseni n. sp. 



Neuropodial 


In segments II and III; 


In segments II, III, and IV; hooks 


hooks: 


hooks more strongly 
bent. 


less strongly bent. 


Notosetae: 


More slender than 


As stout as or stouter than stout- 




stoutest neurosetae; 


est neurosetae; smooth (with- 




with spinous rows 


out spinous rows ) and with en- 




and bifid tips. 


tire tips. 


Notopodia: 


Short, confined to 


Longer, extending to near distal 




middle of neuro- 


tips of neuropodia. 




podial lobes. 




Body width: 


2 mm 


3-4 mm 



The strong neuropodial hooks in the anterior few segments of Austral- 
augeneria certainly suggest an adaptation to some type of commensal 
relationship, such as an association with branched alcyonarians and 
gorgonians. 

Polynoe longicirrus Potts (1910), from the Indian Ocean and taken 
"off a Gorgonian," has been referred to Scalisetosus by Hartman (1959, 
p. 108), perhaps following a suggestion by Augener (1922b, p. 10; foot- 
note ) that it might be a Scalisetosus-hke form; the species may very well 
prove to belong to Australaugeneria. The type-specimens, deposited in 
the British Museum (Natural History), need to be re-examined. As 
pointed out by Hartman (1959, pp. 103, 108), Polynoe longicirrus Potts 
(1910) is a junior homonym of Polynoe (Lepidonotus) longicirra 
Schmarda (1861). 

An attempt was made to re-examine types of Scalisetosus acutipinnis 
Ehlers (1920) from Amboina. However, the four syntypes, deposited in 
the Zoological Museum Berlin, are now completely dry and unsatisfac- 
tory, according to Dr. G. Hartwich ( in htt. ) . 

KEY TO THE SIX GENERA FORMERLY 

REFERRED TO SCALISETOSUS MCINTOSH 

1. Neurosetae of segments II and III modified in the form of stout 

hooks, associated with enlarged presetal hoodhke neuropodial 

lobes. [Body short, about 40 segments. Elytra 15 pairs, on segments 



26 Proceedings of the Biological Society of Washington 

2, 4, 5, 7 . . . 23, 26, 29, 32. Ceratophores of lateral antennae in- 
serted ventrally. Tentacular segment (I) achaetous. Buccal seg- 
ment (II) without nuchal fold. Notosetae stouter than or more 
slender than neurosetae, smooth or with spines, Neuropodia with 
presetal and postsetal lobes rounded, subequal, without projecting 
presetal acicular lobes. Neurosetae without basal semilunar pockets 
or cusps, of 2 or more types, with wide basal regions and long slen- 
der spinous regions and blunt tips (upper ones) and short, stout, 
slightly hooked tips (lower ones). Dorsal tubercles inconspicuous. 

Nephridial papillae distinct on posterior half of body.] 

Australaugeneria new genus 

1'. Segments II and III without neuropodial hooks and modified lobes 2 
2. Neurosetae without basal semilunar pockets or cusps. Tentacular 
segment ( I ) with few setae. Dorsal tubercles nodular. [Body short, 
about 40 segments. Elytra 16 pairs, on segments 2, 4, 5, 7 . . . 23, 
26, 29, 32, 35. Ceratophores of lateral antennae inserted ventrally. 
Buccal segment (II) without nuchal fold. Notosetae much stouter 
than neurosetae, without spinous rows but with few stout spines. 
Neurosetae slender, with faint spinous regions, with tips shghtly 
falcate, entire (lower ones) and minutely bifid (upper ones). 
Neuropodia with long projecting presetal acicular lobes and short 

rounded postsetal lobes. Nephridial papillate distinct.] 

Scalisetosus Mcintosh 



Neurosetae with basal semilunar pockets or cusps. Tentacular seg- 
ment (I) achaetous. Dorsal tubercles inconspicuous 3 

Neuropodia with presetal and postsetal lobes rounded, subequal, 
without projecting presetal acicular lobes. Ceratophores of lateral 
antennae inserted terminally, on same level as median antenna. 
Elytra numerous (28-54 pairs or more), on segments 2, 4, 5, 7 . . . 
23, 26, 29, 32, 33, 35, alternate segments to end of body. [Body 
elongate, 50-110 or more segments. Buccal segment (II) with bul- 
bous area middorsally. Notosetae more slender than neurosetae, 
tapering to blunt tips, very finely serrated. Neurosetae all similar, 
finely serrated, tips slightly to distinctly hooked, entire. Without 
nephridial papillae but with rounded plateHke structures over 

nephridial areas.] Pottsiscalisetosus new genus 

Neuropodia with projecting presetal acicular lobes and short, 
rounded postsetal lobes. Ceratophores of lateral antennae inserted 
ventrally. Elytra 15 (or 16) pairs, on segments 2, 4, 5, 7 . . . 23, 26, 
29, 32 (and sometimes 34), leaving variable number of posterior 

segments without elytra 4 

Body elongate, up to 74 segments, with long posterior region (up to 
42 segments ) lacking elytra. Nephridial papillae indistinct but with 
inflated nephridial areas. [Buccal segment (II) without nuchal 
fold. Notosetae as stout as or stouter than neurosetae, nearly smooth, 
with scattered closely appressed spinous rows along convex border. 
Neuropodial presetal lobes broad, diagonally truncate. Neurosetae 



Review of Scalisetosus 27 

all similar, with long finely spinous regions, tips slightly hooked, 
minutely bidentate.] Adyte Saint-Joseph, EMENDED 

4'. Body short, about 40 segments, with short posterior region ( 12 or 
less segments) lacking elytra. Nephridial papillae distinct 5 

5. Notosetae stouter than neurosetae, curved, sabre-hke, smooth except 
for few spines along convex margin. Neurosetae of two kinds: supra- 
acicular slender, with elongate spinous areas, tips slightly hooked, 
bifid; subacicular stouter, smooth, tips falcate, entire. Neuropodial 
presetal lobes diagonally truncate, subtriangular. Buccal segment 
without nuchal fold . Paradyte new genus 

5'. Notosetae subequal to neurosetae in width, with spinous pouches 
and blunt tips. Neurosetae all similar (lower ones shorter), with 
distal spinous regions, tips hooked, bifid. Neuropodial presetal lobes 
conical. Buccal segment with small nuchal fold _ Subadyte new genus 

Literature Cited 

Alaejos Y. Sanz, L. 1905. Estudio descriptivo de algunas especies 
de PoHnoinos de las costas de Santander. Mem. Soc. Espan. 
Hist. Nat. Madrid, 3: 1-76, 12 pis. 

Augener, H. 1913. Ergebnisse der Hamburger Siidwest-austrahschen 
Forschungsreise 1905. Polychaeta I, Errantia. Die Fauna 
Siidwest-Austrahens, 4 (5): 65-304, 42 figs., 2 pis. 

. 1922a. Results of Dr. E. Mjoberg's Swedish Scientific Ex- 
peditions to Australia 1910^13. Polychaeten. K. Svenska 
Vetensk. Akad. Handl., 63 (6): 1-49, 10 figs. 

. 1922b. Revision der australischen Polychaeten-Typen von 

Kinberg. Ark. Zool. Stockholm, 14 (8): 1-42, 10 figs. 

Benham, W. B. 1915. Report on the Polychaeta obtained by the F.I.S. 
Endeavour on the coasts of New South Wales, Victoria, Tas- 
mania and South AustraHa. Part. I. Sydney, H. C. Dannevig, 
3 (4): 171-237, pis. 38-45. 

Claparede, fi. 1868. Les AnneHdes Chetopodes du Golfe de Naples. 
Mem. Soc. Phys. Hist. Nat. Geneve, 19: 313-584, 16 pis. 

. 1870. Les Annehdes Chetopodes du Golfe de Naples. Sup- 
plement. Mem. Soc. Phys. Hist. Nat. Geneve, 20 (2): 365- 
452, 14 pis. 

Day, J. H. 1962. Polychaeta from several localities in the western 
Indian Ocean. Proc. Zool. Soc. London, 139 (4): 627-656, 
5 figs. 

. 1967. A monograph on the Polychaeta of Southern Africa. 

Part 1. Errantia. Publ. Brit. Mus. (Nat. Hist.), London, No. 
656: 1-458, 108 figs. 

Ehlers, E. 1864. Die Borstenwiirmer (Annelida Chaetopoda), 1: 
1-268, 11 pis. 

. 1920. Polychaeten von Java und Amboina. Ein Beitrag 

zur Kenntnis der malaiischen Strandfauna. Abh. K. Ges. 
wiss. Gottingen, N.F., 10 (7): 1-73, 3 pis. 



28 Proceedings of the Biological Society of Washington 

Fauvel, p. 1914. Annelides polychetes non pelagiques provenant des 

Campagnes de YHirondelle et de la Princesse-Alice (1885- 

1910). Res. Camp. Sci. Monaco, 46: 1-432, 31 pis. 
. 1923. Polychetes errantes. Faune de France, 5: 1-488, 181 

figs. 
. 1953. Annelida Polychaeta. The Fauna of India including 

Pakistan, Ceylon, Burma and Malaya. Allahabad, The Indian 

Press, 1-507, 250 figs. 
Grube, E. 1878. Annulata Semperiana. Mem. Acad. Imp. Sci. St. 

Petersbourg, (7), 25 (8): 1-300, 15 pis. 
Hartman, O. 1959. Catalogue of the Polychaetous annelids of the 

World. Allan Hancock Found. Publ. Occas. Paper, No. 23: 

1-628. 
HoRST, R. 1915. On new and little-known species of Polynoinae from 

the Netherlands' East Indies. Zool. Meded. Leyden, 1: 2-20. 
. 1917. Polychaeta errantia of the Sifcoga-Expedition. Pt. 2. 

Aphroditidae and Chrysopetalidae. Siboga-Exped. Leyden, 

24b: 1-140, 5 figs, pis. 11-29. 
Imajima, M. and Hartman, O. 1964. The polychaetous annelids of 

Japan. Part. I. Allan Hancock Found. Publ. Occas. Paper, No. 

26: 1-237, 35 pis. 
IzuKA, A. 1912. Errantiate Polychaeta of Japan. J. College Sci. Imp. 

Univ. Tokyo, 30 (2): 1-262, 24 pis. 
Johnston, G. 1865. A catalogue of the British non-parasitical worms 

in the collection of the British Museum, London, 1-366, 20 

pis. 
Mcintosh, W. C. 1874. On the Invertebrate Marine Fauna and Fishes 

of St. Andrews. Annulata, Discophora, Oligochaeta and 

Polychaeta. Ann. Mag. Nat. Hist. London, ser. 4, 14: 192- 

207. 
. 1876. On British Annelida. Pt. 1. Trans. Zool. Soc. Lon- 
don, 9: 371-394, pis. 67-70. 
. 1885. Annelida Polychaeta. In, Report on the scientific re- 
sults of the voyage of H.M.S. Challenger . . . 1873-76 . . . 

Zoology, 12 (34): 1-554, pis. 1-55, 1A-39A. 
. 1900. A monograph of the British Annelids, Pt. 2. Poly- 
chaeta. Amphinomidae to Sigalionidae, 215-442, pis. 24-42. 
Marenzeller, E. von. 1902. Siidjapanische AnneHden. III. Aphro- 

ditea, Eunicea. Denkschr. Akad. Wiss. Wien, 72: 563-582, 3 

pis. 
Monro, C. C. A. 1928. On some Polychaeta of the Family Polynoidae 

from Tahiti and the Marquesas. Ann. Mag. Nat. Hist., ser. 

10, 2: 467-472, 4 figs. 
. 1930. Polychaeta worms. Discovery Reports, 2: 1-222, 

91 figs. 



Review of Scalisetosus 29 

Moore, J. P. 1903. Polychaeta from the coastal slope of Japan, and 
from Kamchatka and Bering Sea. Proc. Acad. Nat. Sci. Phila- 
delphia, 55: 401-490, pis. 23-27. 

Okuda, S. 1936. Japanese commensal polynoids. Annot. Zool. Japon., 
15: 561-571, 7 figs. 

Potts, F. A. 1910. Polychaeta of the Indian Ocean. Pt. 2. The Palmy- 
ridae, Aphroditidae, Polynoidae, Acoetidae and SigaUonidae. 
Trans. Linn. Soc. Zool., ser. 2, 16: 325-353, pis. 18-21. 

Saint- Joseph, Baron de. 1899. Annelides polychetes de la rade de 
Brest et Paimpel. Ann. Sci. Nat. Paris, ser. 8, 10: 161-194, 
pi. 6. 

ScHMARDA, L. K. 1861. Neue wirbellose Thiere beobachtet und ges- 
ammelt auf einer Reise um dei Erde 1853 bis 1857. Leipzig, 
vol. 1. Turbellarien, Rotatorien und Annehden. Pt. 2, 1-164, 
22 pis. 100 figs. 

St0p-Bowitz, C. 1948. Polychaeta from the Michael Sars North At- 
lantic Deep-Sea Expedition 1910. Rep. Sci. Results Michael 
Sars N. Atlantic Deep-Sea Exped. 1910, 5 (8): 1-91, 51 
figs. 

WiLLEY, A. 1905. Report on the Polychaeta collected by Professor 
Herdman, at Ceylon, in 1902. Ceylon Pearl Oyster Fisheries, 
Supp. Rep. Pt. 4, no. 30: 243-324, 8 pis. 



30 Proceedings of the Biological Society of Washington 



QH 

1 

B4X 



82, pp. 31-42 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



REMARKS ON THE NORTH PACIFIC HARMOTHOE 

TENEBRICOSA MOORE (POLYCHAETA, POLY- 

NOIDAE) AND ITS ASSOCIATION WITH 

ASTEROIDS (ECHINODERMATA, ASTEROIDEA). 

By Marian H. Pettdbone 
Smithsonian Institution, Washington, D. C. 

Ectocommensal associations between various animal groups 
are often overlooked during routine trawling or dredging op- 
erations unless special attention is given to removing the com- 
mensal animals from their hosts at the time the collections are 
made. For example, in the usual method of sorting, a polynoid 
commensal would be preserved with the echinoderm host or, 
should it become detached, it would remain free in the trawl 
material and be sorted with the other polychaetes, with no in- 
dication of its association. The receipt of a sizeable collection of 
polynoid worms, observed to have been ectocommensal on 
asteroids, initiated the preparation of this report. This collec- 
tion was made by Mr. Miles S. Alton during his study of the 
bathymetric distribution of the Asteroidea off the northern 
Oregon coast, in trawls made by the M/V Commando and 
John N, Cobb in 1961 to 1964 (Alton, 1966; fig. 1). The poly- 
noids were found associated with three species of sea stars, 
two species of the family Solasteridae, Solaster borealis (Fisher) 
and Heterozonias alternatus (Fisher), and one species of the 
family Goniasteridae, Hippasteria californica Fisher. Some 172 
polynoid specimens were collected from 17 stations (32 col- 
lections ) in 165 to 1050 fathoms. Of these, 120 specimens were 
taken from H. alternatus, 27 from S. borealis, and 4 from H. 
californica (see Table I, columns 4-6). In addition, 21 speci- 
mens were found free in the trawl material and sorted with 
the other polychaetes; they were probably removed acciden- 

2^Proc. Biol. Soc. Wash., Vol. 82, 1969 (31) 



,^THS0/v7^^ 

JUN 2 1969 



32 Proceedings of the Biological Society of Washington 



I 



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Harmothoe tenebricosa 



33 



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Fig. 1. U. S. Bureau of Commercial Fisheries- Atomic Energy Com- 
mission trackline study area off the northern Oregon coast, showing 
approximate trawHng station locations of M/V Commando and John N. 
Cobb, 1961-1964, from 50 to 1050 fathoms (91 to 1920 meters). (Pre- 
pared by Miles S. Alton. ) 



tally from their sea star hosts. That this is a definite possibility 
is indicated by the fact that all 21 specimens were obtaiaed 
from stations where at least one of the three sea star hosts was 
also taken (Table I, columns 4-7). In addition to the collection 
of polynoids from off Oregon, three specimens of H. tenebri- 
cosa were collected in the Bering Sea in 250 fathoms by the 
M/V Arthur H., where they were associated with Solaster 
borealis. 

The polynoids were identified as Harmothos tenebricosa 
Moore, originally described from three specimens collected 
by the Steamer Albatross off Southern California in 500 to 800 
fathoms (Moore, 1910). The same species was later described 
by Treadwell, as Eunoe exoculata, from two specimens col- 
lected by the Albatross from Lower California in 475 fathoms 
(Treadwell, 1923) and subsequently referred to H. tenebri- 
cosa by Hartman (1938, p. 118). Uschakov (1950) described 



SWiiHSONiAK MAY 2 9 1969 



34 Proceedings of the Biological Society of Washington 



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36 Proceedings of the Biological Society of Washington 

Harmothoe pellucelytris from eleven specimens collected by 
the R/V Vityaz in the Okhotsk Sea in 232 to 328 fathoms; H. 
pellucelytris is herein designated a synonym of H. tenehricosa. 
Levenstein (1961) reported the species (as H. pellucelytris) 
from nine Vityaz stations in the deep parts of the Bering Sea 
in 422 to 1088 fathoms. Berkeley (1966) recorded three speci- 
mens of H. tenehricosa from British Columbia in 505 and 800 
fathoms. In none of the above five records was the association 
of the polynoids with asteroids reported. On the catalogue card 
of the paratype of H. tenehricosa (USNM 17153), collected 
from Alhatross station 4528, is written the note: "Commensal 
with salmon-colored Solaster"; this note was somehow over- 
looked by Moore when the species was described and by Hart- 
man (1938, p. 118) when the types were re-examined. Fisher 
(1911, p. 323) recorded two specimens of Solaster horealis 
from this same Alhatross station. The holotype of H. tenehri- 
cosa (USNM 16877) was collected at Alhatross station 4400 
from which Fisher (1911, p. 333) recorded three specimens of 
Meter ozonias alternatus. The records of H. tenehricosa (includ- 
ing the synonyms Eunoe exoculata and Harmothoe pellucelyt- 
ris) are for the most part within the known geographic and 
bathymetric distributions of the three sea star hosts. See Ta- 
ble 11. 

The locations of the stations off northern Oregon at which 
the commensal polynoid, Harmothoe tenehricosa, were taken 
are shown on Figure 1 and listed in Table I. The collections 
were made in 50 to 1050 fathoms (91 to 1920 meters) during 
the U. S. Bureau of Commercial Fisheries- Atomic Energy 
Commission trawling investigations in 1961 to 1964. For fur- 
ther details of this study see Alton (1966). 

This study was aided in part by a grant from the National 
Science Foundation (GB-1269). I am grateful to Mr. Miles 
Alton for the polychaete material on which this work was 
based, for providing the station data, for preparing the map 
of the collecting area (Fig. 1), for recording the associations 
of the polynoids with the asteroids, for the identifications of 
the latter, and for reading my preUminary manuscript. The 
manuscript benefited from the suggestions of H. H. Hobbs, Jr. 
and D. L. Pawson, both of the Smithsonian Institution. 



Harmothoe tenebricosa 37 

FAMILY POLYNOIDAE MALMGREN 

Harmothoe tenebricosa Moore 

Fig. 2 

Harmothoe tenebricosa Moore, 1910, p. 351, pi. 29, figs. 23-28.— Hart- 
man, 1938, p. 118.— Berkeley, 1966, p. 841.— Alton, 1966, p. 1705. 

Eunoe exoculata Treadwell, 1923, p. 4, figs. 1-4. 

Harmothoe pellucelytris Uschakov, 1950, p. 161, fig. 5, pi. 1, fig. 2; 
1955, p. 154, fig. 39; 1965, p. 135, fig. 39.— Levenstein, 1961, p. 152. 

Material examined: Albatross station 4400, north of San Diego, Cali- 
fornia, 32° 50'N, 118" 03' W, 500-507 fathoms, green mud, 1904.— Holo- 
type of H. tenebricosa ( USNM 16877 ) . Albatross station 4528, Monterey 
Bay, Point Pinos Light, 545-800 fathoms, soft grey mud, 1904. — Para- 
type of H. tenebricosa (USNM 17153). Albatross station D5698, off 
Point Surf, Lower California, 475 fathoms, 1911. — ^Paratype of Eunoe 
exoculata Treadwell (USNM 19148). 

International Pacific Halibut Commission, M/V Arthur H., Bering 
Sea, 54° 30' N, 166° W, 250 fathoms, June 4, 1963, on Solaster borealis 
—3 specimens (USNM 34051). 

U. S. Bureau of Commercial Fisheries-Atomic Energy Commission, 
M/V Commando, M/V John N. Cobb, off northern Oregon coast, 165- 
1050 fathoms, mud, silty-clay, clayey-silt, on asteroids Solaster borealis, 
Heterozonias alternatus, Hippasteria californica, and free in trawl hauls, 
M. S. Alton, collector, 1961-1964—172 specimens (USNM 34001-34006, 
34014-34032, 34039-34050, 34052-34053). 

Description: Length 17-48 mm, width including setae 5 to 16 mm, 
segments 31 to 43. Body flattened dorsoventrally, widest in middle re- 
gion, tapering gradually anteriorly and more so posteriorly. Elytra and 
elytriphores 14-15 pairs, on segments 2, 4, 5, 7, alternate segments to 
23, 26, 29, and 32. Anterior pair of elytra (fig. 2b) small, circular; rest 
of elytra larger, subreniform, overlapping and nearly covering dorsum 
(fig. 2c); elytra soft, membranous, colorless, lacking marginal papillae 
and tubercles except for numerous colorless microtubercles on anterior 
curved part where covered by more anterior overlapping elytra; some 
elytra with elongate-conical papillae near posterior border. 

Prostomium (fig. 2a) bilobed, slightly wider than long, with distinct 
cephalic peaks; 4 eyes small, of pale color or completely lacking pigment. 
Median antenna with short cyhndrical ceratophore in frontal fissure of 
prostomium; style long, slender, tapering gradually to filamentous tip. 
Lateral antenna with short ceratophores, inserted ventrally; styles about 
half length of median antenna. Palps longer than median antenna, ta- 
pered, with short filamentous tips. Tentacular segment (I) with 2 pairs 
tentacular cirri lateral to prostomium, dorsal pair similar to median an- 
tenna, ventral pair shorter; cirrophore with single seta; facial tubercle 
sHghtly developed ventrally. Buccal segment (II) with first pair of 
elytra; ventral or buccal cirri similar to lower tentacular cirri. 

Parapodia subbiramous, prominent, as long as or longer than body 



38 Proceedings of the Biological Society of Washington 




Fig. 2. Harmothoe tenebricosa: a, Anterior end, dorsal view; elytra 
omitted; only bases of palps and tentacular cirri shown; b, first 
elytron; c, elytron from middle of body; d, style of dorsal cirrus; e, ely- 
trigerous parapodium, anterior view; f, cirrigerous parapodium, posterior 
view, style of dorsal cirrus omitted; g, notoseta; h, neuroseta. 



Harmothoe tenebricosa 39 

width, with acicula dark basally, tapering distally to slender colorless 
tips projecting slightly from rami; setae yellow amber-colored, sHghtly 
darker distally (fig. 2e, f). Notopodia low, rounded, with lower part 
prolonged into digitiform acicular process. Notosetae moderate in num- 
ber, rather short, forming spreading tufts. Notosetae slightly stouter than 
neurosetae, straight, nearly smooth except for indistinct spinous rows 
along one border, tapered to blunt tips (fig. 2g). Neuropodia with neuro- 
setae arranged in vertical series; presetal lobes diagonally truncate, 
upper parts shghtly projecting as rounded supra-acicular process; post- 
setal lobes shorter, rounded. Neurosetae longer and more slender than 
notosetae, with slightly enlarged distal spinous regions; transverse spinous 
rows low, close and numerous; tips hooked, with slender subterminal 
accessory tooth; latter smaller or absent in lower neurosetae (fig. 2h). 
Dorsal cirri with elongate cylindrical cirrophores; styles extending be- 
yond neurosetae, slender, tapering to filamentous tips (fig. 2d). Ventral 
cirri shorter than neuropodial lobes, tapering to slender tips. Dorsal 
tubercles, medial to dorsal cirriphores, subcorneal. Ciliated areas on 
anterior and posterior sides of parapodia, elytriphores and dorsal tuber- 
cles; transverse ciHated bands extending between elytriphores and dorsal 
tubercles, two bands per segment. Nephridial papillae beginning on 
segment VI, continuing posteriorly, and extending dorsally between 
parapodia. 

Pygidium with pair of long slender anal cirri, similar to dorsal cirri. 
Pharynx (not extended in available specimens) large, muscular, some- 
what reddish, extending to segment 14, causing body to bulge some- 
what in this region, sometimes visible through thinner middorsal body 
wall; 4 amber-colored jaws, darker distally. 

Color ( preserved ) : whitish, yellowish to brownish, without pigmented 
pattern; elytra colorless. 

Biology: Fragmentation and regeneration appear to be rather com- 
mon in Harmothoe tenebricosa as indicated by the presence of regenerat- 
ing posterior ends on some individuals, as well as smaller regenerating 
elytra, antennae, and dorsal cirri. Some specimens of H. tenebricosa, 
collected in 600 to 900 fathoms on May 27, 29, and 31, 1964, were 
filled with small eggs (approximately 87 ^ in greatest diameter), with 
large germinal vesicles. In one of the collections from station 17A, 
numerous small specimens were collected in 450 fathoms on August 27, 
1963; some were only 10 mm in length, with 31 segments and 14 pairs of 
elytra. In this same collection, 28 specimens of H. tenebricosa were col- 
lected from 19 specimens of Heterozonias alternatus, indicating that some 
sea stars serve as hosts to at least two of the polynoids, at least when 
the polynoids are young. 

As indicated above and on Tables I and II, Harmothoe tenebricosa has 
been found associated with three species of sea stars, Solaster borealis, 
Heterozonias alternatus, and Hippasteria californica. According to Alton 
(1966, p. 1705), the polynoids were found on the oral sides of the 
discs of the sea stars, usually on the interradial areas but occasionally 



40 Proceedings of the Biological Society of Washington 

within the ambulacral grooves. All three species of sea stars are con- 
fined to the North Pacific and commonly occur in the lower bathyal zone 
(275-500 fathoms) and bathyal-abyssal transitional zone (585-850 
fathoms ) . H. alternatus, which bore the largest number of polynoid com- 
mensals (120 specimens), is one of the most common sea stars in the 
lower bathyal zone and the dominant sea star in the bathyal-abyssal 
transitional zone but is far less important at greater depths (Fisher, 1911; 
Alton, 1966; Table II). Hippasteria calif ornica has essentially the same 
bathymetric distribution as Heterozonias alternatus but is less common; 
it was infested by the fewest number of commensal polynoids (4 speci- 
mens) (Fisher, 1911; Alton, 1966; Table II). Both species of sea stars 
have approximately the same geographic distribution — from Washing- 
ton to southern California. 

Solaster borealis harbored less than a fifth as many polynoid com- 
mensals as did H. alternatus (27 specimens). S. borealis has the most 
extensive geographic distribution of the three species of sea stars — 
from Japan, Okhotsk Sea, Bering Sea to Southern Cahfomia, from 60 to 
1222 fathoms and probably beyond, and forms an important part of the 
sea star fauna in the lower bathyal and bathyal-abyssal transitional zones 
(Fisher, 1911; Hayashi, 1940; Djakonov, 1950; Baranova, 1957; Alton, 
1966; Table II). Similarly, the commensal polynoid, Harmothoe tenebri- 
cosa, has approximately the same range — Okhotsk Sea, Bering Sea to 
Lower Cahfomia, in 165 to 1088 fathoms (Table II). 

Distribution: Okhotsk Sea, Bering Sea to Lower Cahfomia. In 165 
to 1088 fathoms. 

Literature Cited 

Alton, M. S. 1966. Bath>Tiietric distribution of sea stars (Asteroidea) 
off the northern Oregon coast. Joum. Fish. Res. Bd. Canada, 
23, (11): 1673-1714, 9 tables, 5 figs. 

Baranova, Z. I. 1957. [Echinoderms of the Bering Sea]. Issled. 
dahievost morei SSSR (Explorations Mers d'USSR), 4: 149- 
266, 19 figs. [In Russian] 

Berkeley, C. 1966. Records of some species of Polychaeta new to 
British Columbia and of extensions in distribution of some 
others. Canadian Joum. Zool., 44: 839-849. 

Djakonov, A. M. 1950 [Sea stars (Asteroidea) of the far-eastern 
seas of the U.S.S.R.]. Akad. Nauk SSSR. OpredehteH po 
Faune SSSR, No. 34: 1-152, 212 figs. [In Russian] 

Fisher, W. K. 1911. Asteroidea of the North Pacific and adjacent 
waters. Bull. U. S. Nat. Mus., No. 76: 1-419, 122 pis. 

Hartman, O. 1938. The types of the polychaete worms of the fami- 
Hes Polynoidae and Polyodontidae in the U. S. National Mu- 
seum and the description of a new genus. Proc. U. S. Nat. 
Mus., 86: 107-134, 41 figs. 

Hayashi, R. 1940. Contributions to the classification of the sea-stars 



Harmothoe tenebricosa 41 

of Japan. I. Spinulosa. Joum. Fac. Sci. Hokkaido Imp. Univ., 

ser. 6, 7: 107-204, pis. 7-13. 
Levenstein, R. J. 1961. [Polychaete worms (Polychaeta) from the 

deep part of the Bering Sea.] Trudy Inst. Okeanol. Akad. 

Nauk SSSR, 46: 147-178, 10 figs. [In Russian] 
Moore, J. P. 1910. The polychaetous annehds dredged by the U.S.S. 

Albatross off the coast of southern California in 1904: II 

Polynoidae, Aphroditidae and Segaleonidae. Proc. Acad. Nat. 

Sci. Philadelphia, 62: 328-402, pis. 28-33. 
Treadwell, a. L. 1923. Polychaetous annelids from Lower California 

with descriptions of new species. Amer. Mus. Novitates, no. 

74: 1-11, 8 figs. 
UscHAKOv, P. V. 1950. [Polychaete worms (Polychaeta) from the 

Okhotsk Sea.] Issled. dalnevost morei SSSR (Explorations 

Mers d'USSR), 2: 140-234, 39 figs., 2 pis. [In Russian] 
. 1955. [Polychaeta of the far-eastern seas of the U.S.S.R.]. 

Akad. Nauk SSSR. Opredeliteli po Faune SSSR, No. 56: 

1-445, 164 figs. [In Russian] 
. 1965. Polychaeta of the far-eastern seas of the U.S.S.R. 

(English translation). Pp. 1-419, 164 figs. Jerusalem Israel 

Program for Scientific Translations. 



42 Proceedings of the Biological Society of Washington 



l^rOC?^ 



Vol. 82, pp. 43-62 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



THE GENERA POLYEUNOA McINTOSH, 

HOLOLEPIDELLA WILLEY, AND THREE NEW 

GENERA (POLYCHAETA, POLYNOIDAE) 

By Marian H. Pettibone 
Smithsonian Institution, Washington, D. C. 

The polynoid genera Polyeunoa Mcintosh (1885) and Holo- 
lepidella Willey (1905) have been used by various polychaete 
workers to include identical species. The species referred to 
the two genera are generally elongate, with numerous segments 
and numerous pairs of elytra. In addition, the lateral antennae 
are inserted ventrally on the prostomium, i. e., harmothoid 
type of insertion. The elytra in the posterior segments may be 
rather irregularly arranged in these elongate species. In a study 
of numerous specimens of the type-species of Polyeunoa, P. 
laevis Mcintosh, Bergstrom (1916) pointed out the unusual 
variability in the number and arrangement of the elytra. At the 
same time, he suggested that Polynoe tuta Grube and Hololepi- 
della commensalis Willey might both be referred to Polyeunoa, 
since these two species also show variability in the arrange- 
ment of the elytra in the posterior regions. The elytral variabil- 
ity was well shown for Polynoe tuta (as Halosydna) by John- 
son (1901). Augener (1918, 1922) suggested that Hololepidella 
be used for polynoids with a harmothoid prostomium and nu- 
merous elytra extending along the body. Neither Bergstrom nor 
Augener emphasized differences in the arrangement of the ely- 
tra in the anterior region or other parapodial differences. Berke- 
ley and Berkeley ( 1948) and Pettibone ( 1953), following Berg- 
strom, referred Polynoe tuta Grube to Polyeunoa Mcintosh; 
emended Bergstrom. The same species was referred to Holo- 
lepidella, following Augener, by Annenkova (1937), Hartman 
(1948), and Uschakov (1955). Similarly, Polynoe nigropunct- 
ata Horst was referred to Hololepidella by Day (1957)-^nd to, 

3— Prog. Biol. Soc. Wash., Vol. 82, 1969 



44 Proceedings of the Biological Society of Washington 

Polyeunoa by Day (1967). Thus, it was more or less generally 
understood that the arrangement of the elytra in these elongate 
species was so variable that their exact arrangement was not 
especially important and, as such, often was not included in 
the original descriptions. 

Recently, Devaney (1967) pointed out some characters of 
Hololepidella that had been overlooked, particularly concern- 
ing the arrangement of the elytra in the anterior region and 
some setal differences. It was of interest to follow this further 
and to re-examine as far as possible the species that have been 
referred to the two genera. Material was available for study 
in the British Museum (Natural History) (BMNH), Zoological 
Museum Amsterdam (ZMA), Zoological Museum Hamburg 
(ZMH), Zoological Museum Berlin (ZMB), and the United 
States National Museum (USNM). While the arrangement of 
the elytra in the posterior regions of these elongate species 
may vary considerably, their disposition in the anterior re- 
gions appears to be constant. Additional setal and parapodial 
differences, especially the shape of the neuropodial lobes, 
are sufficient to separate the species herein reviewed into 
five genera, three of which are new. The study is summarized 
by providing a key to these five polynoid genera. 

The following genera and species are reviewed (species for 
which specimens were available for study are indicated by an 
asterisk): 

Polyeunoa Mcintosh; *P. laevis Mcintosh 

emended Bergstrom p. flynni (Benham) new com- 

bination 
P. dubia Hartmann-Schroder 
Hololepidella Willey H. commensalis Willey 

H. venosa (Grube) new combi- 
nation 
^H.nigropunctata (Horst) 

Neohololepidella new genus *N. murrayi new species 

( =:Hololepidella commensalis. — 
Monro, 1937; Not Willey, 
1905) 



Islew polynoid polychaetes 45 

Paraholohpidella new genus "^P. greeffi (Augener) new com- 
bination ( includes Hololepidella 
fagei Rullier) 

Grubeopolynoe new genus "^G.tuta (Grube) new combina- 
tion 
G. semenovi ( Annenkova ) new 
combination 

This study was aided in part by a grant from the National 
Science Foundation (GB-1269). For the loan of the speci- 
mens, I wish to extend thanks to R. Sims of the British Mu- 
seum (Natural History), to S. van der Spoel of the Zoological 
Museum Amsterdam, to G. Hartmann-Schroder of the Zoo- 
logical Museum Hamburg, and to G. Hartwich of the Zoo- 
logical Museum Berlin. The manuscript benefited from the 
suggestions of F. A. Chace, Jr. and M. L. Jones, both of the 
Smithsonian Institution. 

FAMILY POLYNOIDAE MALMGREN 

Genus Polyeunoa Mcintosh, 1885; EMENDED Bergstrom, 1916 

Type-species: P. laevis Mcintosh, 1885, by monotypy. Gender: fem- 
inine. 

Diagnosis: Body long, flattened, widest anteriorly, tapered poste- 
riorly, segments numerous ( up to 100 ) . Elytra at least 15 pairs, arranged 
on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 32, with or with- 
out some additional elytra sporadically and irregularly arranged (1 to 
14 or more extra pairs; may be asymmetrical). Elytra rather large in an- 
terior region, nearly covering dorsum; small in posterior region, when 
present; without fringes of papillae or tubercles. Prostomium bilobed, 
with cephalic peaks distinct, poorly developed, or absent, with 2 palps 
and 3 antennae; ceratophore of median antenna in anterior notch of 
prostomium; lateral antennae with distinct ceratophores, inserted ven- 
trally below level of median antenna. Tentacular segment (I) with 2 
pairs tentacular cirri, without setae; without conical facial tubercle. Buc- 
cal segment (II) with long ventral buccal cirri; without nuchal fold. 
Parapodia subbiramous. Notopodia forming well-developed digitiform 
acicular lobes; notosetae stout (as stout as or stouter than neurosetae), 
short to long, smooth or very lightly striated, tapering to blunt tips. 
Neuropodia with prominent conical presetal acicular lobes and shorter 
rounded postsetal lobes; neurosetae stout, with enlarged spinous regions 
and bare, nearly straight, entire tips (may be weak indications of sec- 
ondary tooth); neurosetae all similar except upper ones slightly stouter; 
neurosetae of buccal segment (II) more slender than those following. 



SS: MAY 2 9 1989 



46 Proceedings of the Biological Society of Washington 

Dorsal cirri with cylindrical cirrophores. Dorsal tubercles inconspicuous. 
Ventral cirri subulate. Nephridial papillae short, cyHndrical. 

Polyeunoa laevis Mcintosh 
Fig.l 

Polyeunoa laevis Mcintosh, 1885, p. 76, pi. 12, fig. 2, pi. 20, fig. 8, pi. 

7A, figs. 12, 13.— Bergstrom, 1916, p. 288, pi. 3, fig. 7.— Hartman, 

1959, pp. 66, 98; 1964, p. 42, pi. 12, figs. 5-7; 1967, p. 38.— Uscha- 

kov, 1962, p. 175.— Day, 1967, p. 54, fig. 1.5, 1-q. 
Enipo rhombigera Ehlers, 1908, p. 47, pi. 4, figs. 1-12. 
Polyeunoa rhombigera. — Hartman, 1967, p. 39. 

Material examined: Challenger station 145A, off Prince Edward Islands, 
46° 41' S, 38° 10' E, 567 meters, volcanic sand, December 27, 1873 — 
10 Syntypes (BMNH 1885: 12: 1: 55); 2 Syntypes (USNM 4834, 
38974). 

U. S. Navy Antarctic Expedition, Marguerite Bay, Antarctica, Feb- 
ruary 19-22, 1948, D. C. Nutt, collector: Stations 189, 231, 236, in 64 
and 73 meters— 3 specimens (USNM 23863, 23864, 23870); 155-192 
meters, on Thouarella (Parathouarella) variabilis — 1 specimen (USNM 
35302). 

Deep Freeze II, USS Staten Island, Weddell Sea, Antarctica, W. H. 
Littlewood, collector: 71° 18' S, 13° 32' W, 238 meters, December 27, 
1956.— 1 specimen; 77° 32' S, 44° 45' W, 283 meters, January 21, 1957 
— 1 specimen. 

Deep Freeze II, USS Northwind, McMurdo Sound, Antarctica, J. Q. 
Tierney, collector. Station 5, 77° 51' S, 166° 37' E, March 11, 1957— 
1 specimen. 

Deep Freeze IV, USS Edisto, Weddell Sea, Antarctica, J. Tyler, col- 
lector: Sta. ED-16, west of Cape Norwegia, 71° 45' S, 15° 36' W, 1271 
meters, January 23, 1959—5 specimens. Sta. ED-18, 71° 40' S, 15° 35' 
W, 1554 meters, January 25, 1959 — 6 specimens. Sta. ED-20, off Duke 
Ernst Bay, 77° 40' S, 30° W, 384 meters, January 28, 1959—1 specimen. 
Sta. ED-21, 77° 40' S, 35° 30' W, 411 meters, January 30, 1959—1 
specimen. 

Deep Freeze IV, USS Edisto, Palmer Peninsula, J. Tyler, collector: 
Sta. ED-28, Victor Hugo Island, 65° 08' S, 66° 04' W, 133 meters, 



Fig. 1. Polyeunoa laevis (a, Paratype, USNM 4834; b-h, USNM 
35302): a, Prostomium, dorsal view; cephalic peak developed on left 
side, absent on right side; b, anterior end, dorsal view; left antenna re- 
generating; left elytron missing; c, prostomium of same, dorsal view; 
d, cirrigerous parapodium, posterior view; only base of dorsal cirrus 
shown; e, elytrigerous parapodium, anterior view; elytron removed; f, 
neuroseta from segment II; g, short and longer notosetae; h, upper, mid- 
dle and lower neurosetae. 



'New polynoid polychaetes 



47 









M 




li 




E 


- 




lO 


n 




o 

A 


TIT 













9 




48 Proceedings of the Biological Society of Washington 

March 22, 1959—1 specimen. Sta. ED-31, Nansen Island, 66° 20' S, 67° 
47' W, 325 meters, March 24, 1959 — 4 specimens. Sta. ED-34, west 
coast Palmer Peninsula, 66° 09' S, 67° 53' W, 410 meters, April 1, 1959 
—1 specimen. Sta. ED-35, Nansen Island, 65° 58' S, 66° 51' W, 154 
meters, April 4, 1959—8 specimens (USNM 35293-35304). 

Description: As in generic diagnosis. One syntype of 66 segments has 
16 pairs of elytra, the extra pair located on segment 45. Another incom- 
plete syntype of 59 segments has an extra pair of elytra on segment 38. 
Another syntype of 75 segments has 3 extra pairs of elytra on segments 
36, 54, and 59. The styles of the median antenna and upper tentacular 
cirri are very long; those of the lateral antennae are short. Eyes are vari- 
able in size, from moderate to rather large, the anterior pair located on 
the middle and widest part of the prostomium. Notosetae are few to 
moderate in number, usually 5-6 (1-15). Neurosetae are moderate in 
number (about 12). The neurosetae of the buccal segment (II) are 
similar to those of the following segments, but more slender. The styles 
of the dorsal cirri are variable in length, some may be exceedingly long. 
The ventral cirri extend to the tips of the neuropodia. The pharynx has 
9 pairs of papillae and 2 pairs of jaws. The pygidium has a pair of long 
anal cirri. Nephridial papillae begin on segment VI. The dorsal surface 
has 2 transverse ciHated bands per segment. 

Distribution: Antarctic and Subantarctic. Moderate to deep. Asso- 
ciated with arborescent alcyonarians and gorgoneans. 

Remarks: Bergstrom (1916), in a detailed study of numerous speci- 
mens of this species obtained by the Swedish South-polar Expedition, 
commented on the extreme variability in number and arrangement of the 
elytra in the posterior regions (posterior to segment 32). 

It is to be noted that Polyeunoa Mcintosh, emended Bergstrom, was 
incorrectly defined by Berkeley and Berkeley ( 1948 ) and Pettibone 
( 1953 ) when used to include Polynoe tuta Grube. This species is herein 
referred to Grubeopolynoe new genus (see below). Day (1967) included 
in Polyeunoa, P. laevis and Polynoe nigropunctata Horst. The latter spe- 
cies is herein referred to Hololepidella, as has been done by Day ( 1957 ) 
(see below). 

Polyeunoa f lynni ( Benham ) new combination 

Hololepidella flynni Benham, 1921, p. 33, pi. 5, figs. 14-20. — Hartman, 
1959, p. 81. 

Remarks: The types of H. flynni from off Tasmania, collected in 2377 
meters, were not available for study. According to Benham, the elytra 
were arranged on segments 2, 4, 5, 7 . . . 23, 26, 29, 32, 35, 38, 41 
(every third segment), 42, followed by an irregular arrangement of 
elytra and dorsal cirri, including alternate segments and 2 or 3 consecu- 
tive segments; some may be asymmetrical. 

Polyeunoa dubia Hartmann-Schroder 
Polyeunoe [sic] dubia Hartmann-Schroder, 1965, p. 69, figs. 13-17. 



'New polynoid polychaetes 49 

Remarks: P. duhia, described from specimens from Chile in 170 and 
190 meters, appears to be close to P. laevis. The elytra were reported 
on alternate segments from 34 on, with some irregularity in more pos- 
terior segments. 

Genus Hololepidella Willey, 1905 
Type-species: H. commensalis Willey, 1905, by monotypy. Gender: 
feminine. 

Diagnosis: Body rather short, flattened, tapering posteriorly, segments 
up to 55 or more. Elytra up to 26 or more pairs, on segments 2, 4, 5, 
7, alternate segments to 23, 26, 29, 31, 34, alternate segments to end of 
body (may be some irregularity in posterior region). Elytra delicate, 
smooth or with scattered microtubercles, without fringes of papillae. 
Prostomium bilobed, with lobes subtriangular anteriorly, without dis- 
tinct cephalic peaks, with 2 palps and 3 antennae. Ceratophore of me- 
dian antenna in anterior notch of prostomium; lateral antennae with dis- 
tinct ceratophores, inserted ventrally below level of median antenna. 
Tentacular segment (I) with 2 pairs tentacular cirri, without setae; with- 
out conical facial tubercle. Buccal segment ( II ) with longer ventral buc- 
cal cirri; without nuchal fold. Parapodia subbiramous. Notopodia with 
well-developed digitiform acicular lobes; notosetae as stout as or stouter 
than neurosetae, nearly smooth, with few widely separated spines, taper- 
ing to blunt tips. Neuropodia with prominent subtriangular presetal 
acicular lobes and shorter rounded postsetal lobes; neurosetae with 
longer to shorter spinous regions, with tips entire or bifid. Dorsal cirri 
with cylindrical cirrophores. Dorsal tubercles nodular. Ventral cirri subu- 
late. Nephridial papillae short, cylindrical. 

Hololepidella commensalis Willey 
Hololepidella commensalis Willey, 1905, p. 251, pi. 1, figs. 17-20. — 
Hartman, 1959, p. 81. 

Remarks: The type-specimen of H. commensalis Willey from Ceylon, 
taken in 20 to 36 meters from the echinoid Clypeaster humilis, apparently 
no longer exists; at least it is not present either in the British Museum 
(Natural History) (R. Sims, in litt.) or in the University of Liverpool 
( R. J. Pumphrey, in litt. ) . Willey indicated that the incomplete anterior 
fragment of 46 segments had 20 symmetrical pairs of elytra, arranged as 
indicated in the generic diagnosis; an additional pair of elytra were lo- 
cated on segments 44 (left side) and 45 (right side); a posterior frag- 
ment had the elytra and dorsal cirri irregularly alternating. The tips of 
the neurosetae were indicated as entire, thus differing from H. nigro- 
punctata (Horst). 

The record of H. commensalis by Augener (1922), based on 2 speci- 
mens from N. W. Ausbalia on the echinoid Peronella lesueuri, is con- 
sidered doubtful, since the subacicular neurosetae were indicated as hav- 
ing bifid tips. The arrangement of the elytra was not indicated. 

The record of H. commensalis by Fauvel (1932), based on 2 speci- 



50 Proceedings of the Biological Society of Washington 

mens from Mergui, is considered doubtful, since the subacicular neuro- 
setae were stated to have a faint subterminal spur. Here again, the ar- 
rangement of the elytra was not indicated. 

Hololepidella venosa ( Grube ) new combination 
Polynoe venosa Grube, 1878, p. 43, pi. 3, fig. 6.— Hartman, 1959, p. 107. 
Remarks: Willey (1905, p. 251) indicated that his species, Hololepi- 
della commensalis, was close to Polynoe venosa from the PhiHppine Is- 
lands. The latter species agrees with Hololepidella in the arrangement 
of the elytra, at least for the anterior 18 pairs. The type-specimen ap- 
parently no longer exists; at least it is not present either in the Zoological 
Museum Hamburg ( G. Hartmann-Schroder, in litt. ) or in the Zoological 
Museum BerHn (G. Hartwich, in htt.). 

Hololepidella nigropunctata (Horst) 

Fig. 2 

Polynoe nigro-punctata Horst, 1915, p. 20; 1917, p. 104, pi. 21, figs. 

15-17. 
Hololepidella nigropunctata. — Day, 1957, p. 65, fig. 1, a-f. — Devaney, 

1967, p. 287, figs. 1-5. 
Polyeunoa nigropunctata. — Day, 1967, p. 54, fig. 1.5, r-u. 

Material examined: Reef at Ambon anchorage. East Indies — Holotype 
(ZMA 1162). Waikild, Oahu, Hawaii, 2 meters, ectocommensal on the 
ophiuroid, Ophiocoma dentata Miiller and Troschel, June 4, 1965, D. M. 
Devaney, collector— 6 specimens (USNM 33620, 33621). 

Description: As in generic diagnosis. The maximum number of seg- 
ments reported is 55, with 26 pairs of elytra. The elytra are moderately 
imbricated and cover the dorsum; they are smooth except for some micro- 
tubercles on anterior part. The prostomial eyes are moderate in size, 
the anterior pair located in the region of greatest prostomial width. The 
notosetae are slightly curved with widely spaced spines along curved 
edge (8-14 spines). The upper neurosetae are more slender, with longer 
spinous regions and slightly bifid tips; the middle neurosetae have dis- 
tinctly bifid tips, the lower neurosetae have shorter spinous regions and 
slightly falcate entire or indistinctly bifid tips. The dorsal cirri are rather 
long and tapered; the ventral cirri are short, not extending beyond the 
tips of the neuropodia. The pharynx has 9 pairs of papillae and 2 pairs 
of jaws. The pygidium bears a pair of long anal cirri. Nephridial papil- 
lae begin on segment VII. The dorsal surface has 2 transverse dorsal 
ciliated bands per segment. 

Distribution: East Indies, East Africa (Inhaca Island), Central Pacific 
( Hawaii, Johnston Island, Eniwetok ) . Intertidal and dredged. Commensal 
with echinoderms (ophiuroids, asteroids). 

Neohololepidella new genus 
Type-species: N. murrayi new species. Gender: feminine. 
Diagnosis: Body elongate, flattened, subrectangular, segments up to 



ISlew polynoid polychaetes 



51 




Fig. 2. Hololepidella nigropunctata (USNM 33620): a, Anterior 
end, dorsal view; stub of left palp only present; elytra missing; b, anterior 
end, prostomium and upper lip, ventral view; c, elytrigerous parapodium, 
anterior view; elytron missing; d, cirrigerous parapodium, posterior view; 
e, notoseta; f, middle neuroseta; g, lower neuroseta. 



100 or more. Elytra up to 50 or more pairs, on segments 2, 4, 5, 7, al- 
ternate segments to 23, 26, 29, 32, 34, alternate segments to end of 
body. Elytra easily detached. Prostomium bilobed, without distinct 
cephalic peaks, with 2 pair palps and 3 antennae. Median antenna with 
ceratophore in anterior notch of prostomium; lateral antennae with dis- 
tinct ceratophores, inserted ventral to median antenna. Tentacular seg- 
ment (I) with 2 pairs tentacular cirri, with few setae; with bulbous 
facial tubercle. Buccal segment (II) with longer ventral buccal cirri; 
without dorsal nuchal fold. Parapodia subbiramous. Notopodia with 
projecting digitiform acicular lobes; notosetae numerous, thicker than 
neurosetae, nearly smooth, tapering to blunt tips. Neuropodia with long 



52 Proceedings of the Biological Society of Washington 

subtriangular presetal acicular lobes and shorter rounded postsetal lobes; 
neurosetae numerous, with numerous spinous rows, with very short bare 
tips, slightly bifid and entire. Dorsal cirri with cylindrical cirrophores. 
Dorsal tubercles nodular. Ventral cirri subulate. Nephridial papillae 
short, cyhndrical. 

Neohololepidella murrayi new species 
Fig. 3 

Hololepidella commensalis. -Monro, 1937, p. 259. Not Hololepidella com- 

mensalis Willey, 1905. 

Material examined: The John Murray Expedition, station 152, Mal- 
dives, 609-915 meters, from central cavity of calcareous sponge — Holo- 
type (BMNH 1937: 9: 2: 29). 

Description: As in generic diagnosis. The holotype consists of 102 
segments, incomplete posteriorly; it is 40 mm in length, 3 mm in width, 
including parapodia, and 5 mm in width, including setae. The styles of 
the antennae, tentacular cirri, most of the dorsal cirri, and all of the 
elytra are missing. The elytra, as indicated by the elytriphores, con- 
sist of 50 symmetrical pairs (for 102 incomplete segments). No eyes are 
visible on the prostomium (perhaps faded). The cirrophores of the ten- 
tacular segment (I) have a digitiform acicular lobe and 4 setae. Both 
the noto- and neuropodia have projecting acicular lobes, those of the 
notopodia being digitiform and subsetal in position, those of the neuro- 
podia being subtriangular and presetal in position; the presetal lobes 
end bluntly in more anterior segments ( fig. 3, b, c ) ; they have a slender 
appendage in the middle and posterior segments (fig. 3, d). The noto- 
setae are numerous, forming spreading bundles; they are stouter than 
the neurosetae, nearly smooth, with only faint spinous rows on one side 
near the blunt tip. The neurosetae are numerous, subequal in size, taper- 
ing to blunt tips; the lower ones have the tips entire, the rest have mi- 
nutely bifid tips; the spinous rows extend nearly to the tips. The extended 
pharynx has 9 pairs of papillae and 2 pairs of chitinous jaws. The body 
has ciHated transverse dorsal bands between the elytriphores and dorsal 
tubercles, 2 per segment. The nephridial papillae are small and turned 
dorsally between the parapodia, making it difficult to determine on which 
segment they begin. 



Fig. 3. Neohololepidella murrayi (Holotype, BMNH 1937: 9: 2: 
29 ) : a. Anterior end, dorsal view; styles of antennae and tentacular 
cirri, elytra, and notopodia of segment II missing; only base of extended 
pharynx shown; b, elytrigerous parapodium from anterior region of body, 
anterior view; elytron missing; c, cirrigerous parapodium from anterior 
region, posterior view; d, cirrigerous parapodium from middle region of 
body, anterior view; style of dorsal cirrus missing; e, notosetae from 
same; f, neuroseta from same. 



ISIew polynoid polychaetes 



53 




54 Proceedings of the Biological Society of Washington 

Distribution: Central Pacific (Maldives). In 609 to 915 meters. As- 
sociated with calcareous sponge. 

Parahololepidella new genus 

Type-species: Hololepidella greeffi Augener, 1918. Gender: feminine. 

Diagnosis: Body long, slender, flattened, with sides nearly parallel, 
tapered posteriorly, with numerous segments ( up to 140 or more ) . Elytra 
numerous pairs, on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 
32, 33, 35, alternate segments to end of body (may be some irregularity 
in posterior region). Elytra very small, smooth, leaving middorsum and 
parapodia uncovered. Prostomium bilobed, subtriangular anteriorly, with 
or without distinct cephaHc peaks, with 2 palps and 3 antennae. Cera- 
tophore of median antenna in anterior notch; ceratophores of lateral an- 
tennae distinct, inserted ventrally. Tentacular segment (I) with 2 pairs 
tentacular cirri, with single seta; with conical facial tubercle. Buccal seg- 
ment ( II ) with long ventral buccal cirri; without nuchal fold. Parapodia 
subbiramous. Notopodia small, conical; notosetae short, stout (not as 
stout as neurosetae), tapering to blunt tips. Neuropodia with longer 
rounded presetal lobes bearing subacicular digitiform processes and 
shorter rounded postsetal lobes; neurosetae stout, with faint spinous 
regions, and sHghtly hooked, entire tips. Dorsal cirri with cylindrical 
cirrophores, bulbous basally. Dorsal tubercles inconspicuous. Ventral 
cirri tapering. Nephridial papillae short, bulbous. 

Parahololepidella greeffi ( Augener ) new combination 
Fig. 4 

Hololepidella greeffi Augener, 1918, p. 148, pi. 2, figs. 22-24, pi. 3, 

fig. 52, text-fig. 9.— Hartman, 1959, p. 81.— Rullier, 1964, p. 130, 

fig. 3. 
Hololepidella fagei RulHer, 1964, p. 132, fig. 4. — Hartman, 1965, p. 9. 

Material examined: Ilha das Rolas, Ilha Sao Thome, Gulf of Guinea, 
West Africa, R. Greeff, collector— 4 Syntypes (ZMH 5692). 

Description: As in generic diagnosis. The 4 syntypes consist of one 
nearly complete specimen of 136 segments, 3 anterior fragments of 26, 
19, and 17 segments, and a middle and a posterior fragment. They are 
rather hard and shrunken. The elytra are remarkable for their small size 
and are located on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 
32, 33, 35 (not on segments 28, 31, 32, 34, 35, as indicated by Augener); 
from segments 35 to 85, they occur on alternate segments, followed by 
dorsal cirri on segments 86 and 87 and elytra on segment 88, alternating 
to end of body (segment 136). The posterior fragment has some asym- 
metrical segments with an elytron and dorsal cirrus on the same segment. 
The prostomium has distinct cephalic peaks and 2 pairs of large eyes. 
Notosetae are few in number (1-5), nearly smooth, with only faint spi- 
nous rows. The neurosetae are stouter than the notosetae, few in number 
(about 7), with faint spinous rows. The styles of the dorsal cirri are 
long, filiform. The pharynx was not extended and was not examined. 



ISlew polynoid polychaetes 



55 




g ^ hi 

Fig. 4. Parahololepidella greeffi (Syntype, ZMH 5692): a, Pro- 
stomium, dorsal view; styles of all appendages missing; b, elytrigerous 
parapodium from middle region of body, anterior view; elytron missing; 
c, cirrigerous parapodium from middle region of body, posterior view; 
style of dorsal cirrus missing; d, notoseta from same; e, upper and middle 
neurosetae from same; f, elytrigerous parapodium from posterior region 
of body, anterior view; g, notoseta from same; h, upper neuroseta from 
same. 



The nephridial papillae begin on segment VI. The surface of the elytra 
and body are covered with scattered angular extraneous particles. 

Distribution: West Africa (Gulf of Guinea), Cape Verde Islands. 

Remarks: Hololepidella fagei RulHer ( 1964 ) appears to be just a young 
specimen of H. greeffi. Rullier reported specimens of H. greeffi occupy- 
ing white mucous tubes, incrusted with sand grains and fragments of 
shells. Perhaps the tubes were formed by some commensal host. When 
comparing H. fagei with other polynoid species, Rullier ( 1964, p. 134 ) 
confused Hololepidella Willey with Hololepida Moore, which are very 
different genera. 



56 Proceedings of the Biological Society of Washington 

Grubeopolynoe new genus 
Type-species: Polynoe tuta Grube, 1855. Gender: feminine. 
Diagnosis: Body long, flattened, tapering gradually posteriorly, with 
numerous segments (up to 100). Elytra numerous pairs (up to 50 or 
more), on segments 2, 4, 5, 7, alternate segments to 23, 26, 29, 32, 35, 
36, 39, 40, continuing to end of body with two pairs elytra alternating 
with two pairs dorsal cirri; usually with some irregularity on segments of 
posterior region. Elytra large, nearly covering dorsum, smooth, without 
fringes of papillae or tubercles. Prostomium bilobed, without distinct 
cephalic peaks, with 2 palps and 3 antennae. Median antenna with cera- 
tophore in anterior notch of prostomium; lateral antennae with distinct 
ceratophores, inserted ventrally. Tentacular segment (I) with 2 pairs 
tentacular cirri, with single seta or seta lacking; without facial tubercle. 
Buccal segment (II) with long ventral buccal cirri; without nuchal fold. 
Parapodia subbiramous. Notopodia with short digitiform acicular lobes; 
notosetae more slender than neurosetae, of 2 types: shorter, stouter, 
tapering to blunt tips; longer, more slender, tapering to slender tips. 
Neuropodia with diagonally truncate presetal lobes, forming longer supra- 
acicular Ups and shorter rounded postsetal lobes; neurosetae all similar 
except upper ones with longer spinous regions; tips entire or with very 
slender secondary tooth or slight indication of one. Neurosetae of seg- 
ment II different from those following, slender, tapering to slender tips; 
some may be bidentate. Dorsal cirri with cylindrical cirrophores, bulbous 
basally. Dorsal tubercles bulbous. Ventral cirri subulate. Nephridial 
papillae rather long, cyHndrical. 

Remarks: The diagnosis of Hololepidella Willey, as given by Annen- 
kova (1937, p. 147) and Uschakov (1955, p. 135) to include Polynoe 
tuta Grube, applies rather to Grubeopolynoe. The same is true for the 
diagnosis of Polyeunoa Mcintosh, emended Bergstrom, as given by Petti- 
bone (1953, p. 54). 

Grubeopolynoe tuta ( Grube ) new combination 
Figs. 5, 6 
Polynoe tuta Grube, 1855, p. 82. 
Harmothoe tuta. — Johnson, 1901, p. 394, pi. 2, figs. 18, 19, pi. 3, figs. 

20-22. 
Hololepidella tutta [sic]. — Annenkova, 1937, p. 147, pi. 3, figs. 15, 16. 
Hololepidella iwfa.— Hartman, 1948, p. 13; 1959, pp. 81, 106.— Uschakov, 

1955, p. 136, fig. 25. 
Polyeunoa tuta. — Berkeley and Berkeley, 1948, p. 19. — Pettibone, 1953, 

p. 54, pi. 27. 

Material examined: Sitka, Alaska, Dr. Bock, collector — Holotype 
(ZMB 1152). 

Alaska Salmon Investigation, 1903, Karluk, Alaska — 1 specimen 
(USNM 18140). Alaska King Crab Investigation, Station D 3-41, Cas- 
tle Bay, off Chignik Bay, 38-95 meters, June 24, 1941 — 1 specimen 
(USNM 21380). 



New polynoid polychaetes 



57 




Fig. 5. Grubeopolynoe tuta (USNM 25233): a, Anterior end, 
dorsal view, elytra removed; b, prostomium, ventral view; c, two para- 
podia from middle region of body, dorsal view, elytron removed; d, noto- 
setae from segment II; e, neuroseta from segment II. 



Gulf of Georgia, Halibut Bank, British Columbia, Albatross stations 
4193 and 4197, 33-164 meters, 1903—2 specimens (USNM 5737, 5738). 
British Columbia, Mudge and Protection Islands, E. and C. Berkeley, 
collectors— 30 specimens (USNM 35601-35603). 

Washington, M. Pettibone, collector: Brown Island, Olga on Orcas 
Island, Shaw Island, San Juan Archipelago, shore. South John's Island, 
25-46 meters. Off Foulweather Bluff, Puget Sound, 46-55 meters. Ed- 
monds, Meadowdale, Ballard, West Seattle (Alki Point), Puget Sound, 
shore— 50 specimens (USNM 25222-25233, 32319). 

Description: As in generic diagnosis. The type-specimen consists of 
an anterior fragment of 45 segments and a posterior fragment of 81 seg- 
ments. The elytra are distributed as indicated in the generic diagnosis; 
they are all symmetrically arranged, with 2 pairs of elytra alternating 
with 2 pairs of dorsal cirri from setiger 35 on, except for an irregular- 
ity occurring on segments 59 and 60 where only one pair of elytra occurs, 
instead of 2 consecutive pairs. On most of the other specimens examined, 



58 Proceedings of the Biological Society of Washington 




Fig. 6. Grubeopolynoe tuta (a-g, USNM 25233; h-i, USNM 25231): 
a, Cirrigerous parapodium from middle region of body, posterior 
view; b, elytrigerous parapodium from middle region, anterior view, 
elytron removed; c, notosetae from same; d, upper and middle 



Islew polynoid polychaetes 59 

there is some departure from this general pattern, beginning usually on 
segments 35 to 40, but occasionally from segments 29 to 63. This irregu- 
larity consists of asymmetry — an elytron and cirrus on the same segment, 
as well as up to 4 consecutive elytra or dorsal cirri. The notosetae form 
radiating bundles and are moderate in number (about 30). The neuro- 
setae are also moderate in number ( about 30 ) , stouter than the notosetae, 
with short to longer spinous regions; the tips are nearly straight or slightly 
falcate, entire or they may have a delicate secondary tooth. The neuro- 
setae of the buccal segment (II) differ from those following; they are 
more slender, tapering to slender tips, which may be entire or bifid 
(fig. 5, e). The styles of the dorsal cirri taper gradually to fine tips, 
with few scattered minute papillae. The pharynx has 9 pairs of papillae 
and 2 pairs of jaws. The nephridial papillae begin on segment VI; on the 
large specimens, they are extra long and may be equal in length to or 
longer than the ventral cirri. They are found intertidally, living com- 
mensally with the terebelhds Thelepus crispus Johnson and Amphitrite 
robusta Johnson. The dredged forms are characteristically smaller and the 
notosetae may be more slender (fig. 6, h). 

Distribution: North Pacific — Alaska to Oregon, North Japan Sea. In- 
tertidal to 384 meters. Associated with other polychaetes (terebelhds). 

Grubeopolynoe semenovi ( Annenkova ) new combination 

Hololepidella semenovi Annenkova, 1937, p. 148, pi. 3, figs. 17-19. — 
Uschakov, 1955, p. 137, fig. 26.— Hartman, 1959, p. 81. 
Distribution: North Japan Sea, in 170 to 210 meters. Associated with 

other polychaetes ( ampharetids ) . 

Key to the Genera Formerly Referred to 
Polyeunoa McIntosh and Hololepidella Willey 

1. Neuropodia with prominent subtriangular presetal acicular lobes 
and shorter rounded postsetal lobes. Notopodia with prominent 
acicular lobes. Notosetae as stout as or stouter than neurosetae, 
nearly smooth, tapering to blunt tips. [Neurosetae with longer to 
shorter spinous regions, with tips entire or bifid] 2 

1'. Neuropodia without subtriangular presetal acicular lobes. Noto- 
podia short, conical. Notosetae more slender than neurosetae. [Body 
elongate, with numerous segments (up to 100 or more)] 4 

2. Body rather short (up to 55 or more segments). First 15 pairs of 
elytra arranged on segments 2, 4, 5, 7, alternate segments to 23, 26, 



neurosetae from same; e, cirrigerous parapodium from posterior region 
of body, posterior view; f, elytrigerous parapodium from posterior region, 
anterior view; g, middle neuroseta from same, showing slight indication 
of secondary tooth; h, notosetae from middle parapodium of dredged 
specimen; i, neuroseta from same, with delicate secondary tooth. 



60 Proceedings of the Biological Society of Washington 

29, 31, then on segments 34, 36, and alternate segments to end of 
body. [Neurosetae with long bare tips. Dorsal tubercles nodular] 

Hololepidella Willey 

2'. Body elongate, with numerous segments (up to 100 or more). 
First 15 pairs of elytra arranged on segments 2, 4, 5, 7, alternate 
segments to 23, 26, 29, 32 3 

3. With or without some additional elytra irregularly distributed pos- 
terior to segment 32, leaving most of posterior region uncovered. 

Neurosetae with long bare tips. Dorsal tubercles inconspicuous 

Polyeunoa Mcintosh 

3'. Numerous pairs elytra arranged on segments 34 and alternate seg- 
ments to end of body. Neurosetae with very short bare tips. Dorsal 
tubercles nodular Neohololepidella new genus 

4. Elytra arranged on segments 35, 36, 39, 40, then continuing with 2 
consecutive elytra alternating with 2 consecutive dorsal cirri 
(usually with some irregularity in posterior segments). Elytra 
large, nearly covering dorsum. Neuropodia with presetal lobes 
longer, diagonally truncate, forming longer supra-acicular hps and 
shorter rounded postsetal lobes. Notosetae moderate in number, 
spinous, of 2 types: shorter, stouter, curved, tapering to blunt tips 
and longer, more slender, tapering to slender tips. Neurosetae with 
distinct spinous regions, tips entire or with slender secondary tooth. 
Dorsal tubercles conical Grubeopolynoe new genus 

4'. Elytra arranged on segments 33, 35, then continuing on alternate 
segments to end of body. Elytra very small, leaving middorsum and 
parapodia uncovered. Neuropodia with presetal lobes longer, 
rounded, with subacicular digitiform processes and short, rounded 
postsetal lobes. Notosetae few in number (1-5), all similar, short, 
stout, with faint spinous rows, tapering to blunt tips. Neurosetae 
with faint spinous rows and slightly hooked entire tips. Dorsal 
tubercles inconspicuous Parahololepidella new genus 

Literature Cited 

Annenkova, N. p. 1937. [The polychaete fauna of the northern part 
of the Japan Sea.] Issled. Morei USSR, Gosud. Gidrol. Inst. 
Leningrad, 23: 139-216, 12 figs., 5 pis. [In Russian, English 
summary] 

Augener, H. 1918. Polychaeta. Beitrage zur Kenntnis des Meeres- 
fauna Westafrikas. Herausgegeben von W. Michaelsen, Ham- 
burg, 2 (2): 67-625, 111 figs., 6 pis. 

. 1922. Results of Dr. E. Mjoberg's Swedish Scientific Ex- 
peditions to Australia 1910-13. Polychaeten. K. Svenska 
Vetensk. Akad. Handl., 63 (6): 1-49, 10 figs. 

Benham, W. B. 1921. Polychaeta. Australian Antarctic Expedition 
1911-14, under the leadership of Sir Douglas Mawson. Sci- 
entific Reports, ser. C. Zool. Bot., 6 (3): 1-128, pis. 5-10. 

Bergstrom, E. 1916. Die Polynoiden des schwedischen Siidpolar- 



Tslew polynoid polychaetes 61 

Expedition 1901-1903. Zool. Bidr. Uppsala, 4: 269-304, 2 
figs., pis. 2-5. 

Berkeley, E. and Berkeley, C. 1948. Annelida. Polychaeta Errantia. 
In, Canadian Pacific Fauna. Fish. Res. Board Canada, To- 
ronto. Pp. 1-100, 160 figs. 

Day, J. H. 1957. The polychaet fanua of South Africa. Part 4. New 
species and records from Natal and Mocambique. Ann. Natal 
Mus., 14 (1): 59-129, 8 figs. 

. 1967. A monograph on the Polychaeta of Southern Africa. 

Part 1. Errantia. Publ. Brit. Mus. (Nat. Hist.) London, No. 
656: 1-458, 108 figs. 

Devaney, D. M. 1967. An ectocommensal polynoid associated with 
Indo-Pacific echinoderms, primarily ophiuroids. Occas. Pap. 
Bemice P. Bishop Mus., 23 (13): 287-304, 5 figs. 

Ehlers, E. 1908. Die Bodensassigen Anneliden aus den Sammlungen 
der Deutschen Tiefsee-Expedition. Wiss. Ergeb. Deutschen 
Tiefsee-Exped. 1898-1899, 16: 1-168, 23 pis. 

Fauvel, p. 1932. Annelida Polychaeta of the Indian Museum, Cal- 
cutta. Mem. Indian Mus., 12 ( 1 ) : 1-262, 40 figs., 9 pis. 

Grube, E. 1855. Beschreibung neuer oder wenig bekannter Annehden. 
Arch. Naturg. Jahrg., 21 (1): 81-136, pis. 3-5. 

. 1878. Annulata Semperiana. Mem. Acad. Imp. Sci. St. 

Petersbourg, (7) 25 (8): 1-300, 15 pis. 

Hartman, O. 1948. The polychaetous annehds of Alaska. Pac. Sci. 
Univ. Hawaii, 2(1): 3-58, 12 figs. 

. 1959. Catalogue of the Polychaetous annehds of the World. 

Allan Hancock Found. Publ. Occas. Paper, No. 23: 1-628. 

. 1964. Polychaeta errantia of Antarctica. Antarctic Re- 
search Series, 3: 1-131, 39 pis. 

. 1965. Catalogue of the Polychaetous annelids of the World. 

Supplement 1960-1965 and Index. Allan Hancock Found. 
Publ. Occas. Paper, No. 23: 1-197. 

. 1967. Polychaetous annehds collected by the USNS Eltanin 

and Staten Island cruises, chiefly from Antarctic Seas. Allan 
Hancock Monographs in Marine Biology, No. 2: 1-387, 51 
pis. 

Hartmann-Schroder, G. 1965. Zur Kenntnis des subhtorals de chileni- 
schen Kuste unter besonderer Beriicksichtigung der Poly- 
chaeten und Ostracoden. Pt. II. Die Polychaeten des sub- 
htorals. Mitt. Hamburg. Zool. Mus. Inst., 62: 59-305, 300 
figs. 

HoRST, R. 1915. On new and httle-known species of Polynoinae from 
the Netherland's East Indies. Zool. Meded. Leyden, 1: 2-20. 

. 1917. Polychaeta errantia of the Sifcoga-Expedition. Pt. 2. 

Aphroditidae and Chrysopetahdae. Siboga-Exped. Leyden, 
24b: 1-140, 5 figs., pis. 11-29. 



62 Proceedings of the Biological Society of Washington 

Johnson, H. P. 1901. The Polychaeta of the Puget Sound region. 

Proc. Boston Soc. Nat. Hist, 29 (18): 381-437, 19 pis. 
McIntosh, W. C. 1885. AnneHda Polychaeta. In, Report on the sci- 
entific results of the voyage of H.M.S. Challenger . . . 1873- 

76 . . . Zoology, 12 (34): 1-554, pis. 1-55, 1A-39A. 
Monro, C. C. A. 1937. Polychaeta. The John Murray Expedition 

1933-34. Scientific Reports, 4(8): 243-321, 28 figs. 
Pettibone, M. H. 1953. Some scale-bearing polychaetes of Puget 

Sound and adjacent waters. Univ. Washington Press, Seattle, 

Pp. 1-89, 4 figs., 40 pis. 
RxiLLiER, F. 1964. Annelides polychetes. Campagne de la "Calypso": 

lies du Vert. Res. Sci. Campagnes "Calypso," 6: 113-218, 

23 figs. 
UscHAKOv, P. V. 1955. [Polychaeta of the far-eastern seas of the 

USSR.] Akad. Nauk SSSR. OpredehteH po Faune SSSR, No. 

56: 1-445, 164 figs. [In Russian] 
. 1962. [Polychaetous annelids of the families Phyllodocidae 

and Aphroditidae from the Antarctic and Subantarctic] In, 

Biol. Res. Soviet Antarctic Exped. (1955-1958). Explor. 

Fauna Sea I (IX) Akad. Nauk SSSR: 129-189, 9 pis. [In 

Russian] 
WiLLEY, A 1905. Report on the Polychaeta collected by Professor 

Herdman, at Ceylon, in 1902. Ceylon Pearl Oyster Fisheries. 

Supp. Rep. Pt. 4, No. 30: 243-324, 8 pis. 



QH 

1 )1. 82, pp. 63-80 29 May, 1969 

B4X 

NH PROCEEDINGS 

OF THE 
BIOLOGICAL SOCIETY OF WASHINGTON 

REVERSAL OF ASYMMETRY OF CHELAE IN CKLkV?k 
WEBER, 1795 (DECAPODA: OXYSTOMATA) 

By Jackson E. Lewis 
Tulane University^ 

Examination of identified specimens of Calappa in the col- 
lections of the Smithsonian Institution indicates that the asym- 
metry of chelae may be reversed completely, presumably when 
both chelipeds are regenerated simultaneously following autot- 
omy, or only partially, when only a single cheHped has been 
regenerated. Of 809 specimens, seven, or a little less than 1 
percent, exhibit complete reversal and two specimens, partial 
reversal. These phenomena occur with equal frequency on 
males and females and occur rarely also on specimens of other 
genera in the Calappinae. Attributes of the affected specimens 
are described in detail and general criteria are presented 
whereby chelae whose asymmetry has been reversed due to 
regeneration can be recognized. 

In each species of the pantropical and warm temperate ma- 
rine crab Calappa, the proximal portions of the chelae are 
normally similar to one another in size, shape, proportions, 
and ornamentation. However, complex morphological features 
on the dactylus and fixed finger of the right chela render the 
chelae and their individual elements quite distinctive. Because 
the right and left chelae exhibit what may be called 'Tianded- 
ness," they are customarily termed major and minor, respec- 
tively (see Fig. 1). 

Recently, Shoup ( 1968 ) described in detail how the descend- 
ing lobular projection on the dactylus and the complementing 
teeth on the propodus of the major chela are used to secure 



1 Present address: Division of Crustacea, Smithsonian Institution, Washington, 
D,C. 20560. 

4— Proc. Biol. Soc. Wash., Vol. 82, 1969 (63) 



^THSO/VT^ 

r JUN2e)t969 j 



64 Proceedings of the Biological Society of Washington 




Fig. 1. Outline drawing of normal chelipeds (chelae, carpi, and 
"wings" of meri ) of Calappa flammea ( Herbst ) , illustrating features and 
terminology cited in text. Elements are extended to their maximum, 
placing their outer surfaces approximately in the plane of the propodus. 
Bar scale represents 10 mm. A. Right major cheliped. B. Left minor 
cheliped. 



food. An elevated tooth near the base of the fixed finger on 
the outer surface of the major propodus constitutes part of 
this apparatus and is usually extremely stout and conspicuous. 
This tooth is diagnostic of all species of Calappa, but published 
descriptions of the genus and its constituent species seldom 
attribute it specifically to either the right or left propodus. 
Alcock (1896: p. 140) and Rathbun (1937: p. 197) mentioned 
only that one propodus bears the tooth, but Barnard (1950: 
p. 347) noted that it is usually the right. Although varying in 
degree of expression, this tooth is also characteristic of the 
other Recent genera of the subfamily Calappinae, Acantho- 
carpus Stimpson, 1871, Cycloes de Haan, 1837, Mursia Demar- 
est, 1823, and Paracyclois Miers, 1886. 

While studying Eocene decapod remains from Florida, Ross, 
Lewis, and Scolaro (1964) encountered a unique right propo- 
dus which resembled the major propodus of a species of Ca- 
lappa in shape and proportions, but which lacked any trace of 
the characteristic propodal tooth. Following a fruitless search of 
the literature and after consulting with workers having access 
to large collections of Recent species of Calappa, the authors 
concluded with some reservations that the fossil represented 
a new genus distinct from, but closely related to, Calappa. They 
realized, however, that the ultimate answer to the validity of 
the new genus rested in a thorough examination of large num- 
bers of preserved specimens of Calappa. 



Asymmetry of Calappa chelae 65 




Fig. 2. Outline drawing of chelipeds of Calappa hepatica (Linne) 
(USNM 33400), illustrating complete and nearly perfect reversal of 
asymmetry. Elements are extended to their maximum, as in Fig. 1. Bar 
scale represents 10 mm. A. Right minor cheliped. Note smaller size rela- 
tive to left cheliped, undefined teeth on wing of merus, and slightly ab- 
normal proportions of propodus. B. Left major chehped. Note poorly 
defined teeth on wing of merus, presence of supradactylar projection, 
and lack of superior proximal dactylar tooth. 

The purpose of this paper is to report the results of such a 
study. A re-evaluation of the fossil genus in the light of these 
results appears in a separate note (Lewis, 1969). 

Observations 

Specifically, the problem under investigation is whether the 
distinctive characters of a minor chela do occur in species of 
Calappa on a right, customarily major, cheliped. In more gen- 
eral terms, is the asymmetry of the chelae ever reversed either 
partially (only one cheliped affected) or completely (both 
cheHpeds affected)? These phenomena have not heretofore 
been discussed in the literature on these animals. 

During the summer of 1967, the writer examined the vari- 
ability of major and minor chelae on all identified specimens 
of Calappa in the alcoholic and dry collections of the Smith- 
sonian Institution in order to determine ( 1 ) if reversal of asym- 
metry does occur; (2) if so, to what extent and under what 
circumstances; and ( 3 ) the variety and frequency of malforma- 
tions which occur within the genus. Of the 809 specimens 
examined, 15 individuals possess conspicuously abnormal cheli- 
peds. Of these, seven exhibit some degree of reversal of asym- 



SWITHSONIAK MAY 2 9^69 



66 Proceedings of the Biological Society of Washington 

metry on both chelipeds, that is, complete reversal ( see Table 
I and Fig. 2); two of seven other individuals exhibit partial 
reversal, and all seven of these specimens possess one com- 
pletely normal cheHped and one abnormal cheliped which 
usually bears a grotesque propodus (see Table II); and one 
specimen shows the effects of injury to the propodus of one 
cheliped during normal growth (see Table II). 

Considering in detail the characteristics of the left chelipeds 
of the specimens listed in Table I, the merus and carpus are 
quite indistinguishable from those of their unreversed counter- 
parts, but in two cases [C. hepatica (Linne, 1758), USNM 
33400 and 93819] the teeth on the wings of the meri are 
weakly delineated, although quite normal in number. Of appre- 
ciably greater signiFicance among these specimens, the left 
(major) propodus usually retains a prominent supradactylar 
projection, which normally occurs only on the minor propodus, 
and the elevated tooth, which characteristically crowns the su- 
perior proximal margin of the major dactylus, is missing en- 
tirely on the left (major) dactylus (compare Figs. 1 and 2), 
as demonstrated by specimens of C. flammea (Herbst, 1794) 
(USNM 71113), C. nUidu Holthuis, 1958 (USNM 103153), 
C. hepatica (USNM 33400), and C. gallus (Herbst, 1803) 
( USNM 48564 ) . Partial exceptions are noted, however, on C. 
hepatica (USNM 93819), where both the supradactylar pro- 
jection and the tooth are present, but the latter is strongly 
reduced, and on C. ? flammea (USNM 100833) and C. gallus 
(USNM 77211), on each of which the tooth and the supra- 
dactylar projection are absent. 

The right chelipeds in this group of specimens are usually 
slightly smaller than the left, and their individual elements 
are dramatically more aberrant than those of the left. The 
teeth on the wings of the meri are universally fewer than 
normal in number and are usually weakly delineated; on one 
specimen of C. hepatica (USNM 33400), they are almost com- 
pletely absent (see Fig. 2). However, the greatest divergence 
from normalcy lies in the shape and proportions of the right 
(minor) propodi and in the ornamentation on their outer sur- 
faces. The propodi are almost as high as or higher than they 



Asymmetry of Calappa chelae 



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70 Proceedings of the Biological Society of Washington 

are long on C. ? flammea (USNM 100833), C. nitida (USNM 
103153), C. hepatica (USNM 33400), and C. gallus (USNM 
48564). Usually, surface ornamentation is sparse and diverges 
to varying degrees from the typical. Tuberculation is gener- 
ally weak, uneven, and random rather than orderly (C. flam- 
mea, USNM 71113; C. ? flammea, USNM 100833; and C. gallus, 
USNM 48564), and granulation, also typically sparse and ran- 
dom, may be lacking almost entirely (C. flammea, USNM 
71113). On C. niiida (USNM 103153), there is little differ- 
ence in size between tubercles and granules; on C. gallus 
(USNM 48564), tubercles are really random, completely hair- 
less, overgrown granules totally unlike the customary diagnos- 
tic laminae; on C. gallus (USNM 77211), the tubercles are 
strong and sharp or distinctly pegHke. In general, surface or- 
namentation on these propodi is conspicuously sparse, and 
large portions are smooth, or nearly so, with occasional ran- 
domly distributed tubercles and patches of granules. Excep- 
tions are the two specimens of C. hepatica ( USNM 33400 and 
93819), on which ornamentation is virtually indistinguishable 
from that on normal propodi; on the former specimen, tuber- 
culation is weaker on the right than on the left propodus, but 
less pronounced variations in strength of ornamentation on 
opposing propodi were observed occasionally among other- 
wise normal specimens of all species of Calappa examined in 
this study. 

The first seven specimens listed in Table II, marked "[RC]," 
indicate that when only one cheliped is abnormal, the affected 
appendage is invariably small and/or grotesque, and, with a 
single partial exception, the propodus is conspicuously aber- 
rant in shape, proportions, and ornamentation. Five of these 
specimens exhibit the characteristics of a minor appendage on 
the left cheliped (C. flammea, USNM 66441; C. ocellata Hol- 
thuis, 1958, USNM 99990; C. sulcata Rathbun, 1898, USNM 
99980, and the female, USNM 103450; and C. hepatica, USNM 
65355). On four, the teeth on the wings of the left meri are 
fewer than normal in number, and, where well delineated, they 
bear peculiar, atypical terminations; the superior crests on the 
left propodi of C. flammea (USNM 66441) and C. ocellata 
(USNM 99990) bear far more than the normal number of 



Asymmetry of Calappa chelae 71 

teeth and the crest on C. hepatica (USNM 65355) is rounded 
and bumpy, lacking completely any trace of teeth; and the 
outer surfaces of the left propodi are generally smooth with or- 
namentation confined to a few, knobby, often random tuber- 
cles. The female C. sulcata (USNM 103450) represents the par- 
tial exception mentioned above. Like the four specimens just 
discussed, its abnormal left cheliped bears a minor chela, but, 
unlike them, the teeth on the wing of the left merus are nor- 
mal; ornamentation on all parts of the left cheliped is almost 
normal also, but all elements are small and in life they were 
evidently atypically colored, the color in alcohol being grayish 
purple, in marked contrast to the cream to ivory-colored ele- 
ments of the right cheliped. 

The remaining two specimens in this group have charac- 
teristics of a miaor appendage on the right cheliped and 
thereby exhibit partial reversal. The wing of the right merus 
on the male C. sulcata (USNM 103450) has five teeth which are 
strongly hooked distally, and that of C. hepatica (USNM 
99134) has only two unusual quadrate teeth. In proportions, 
relative size, and ornamentation, the right propodi of these 
specimens resemble the strongly aberrant left propodi on the 
four other members of this group discussed earlier. 

The only abnormality observed during this study which was 
caused solely by injury during normal growth occurs on C. 
hepatica (USNM 65418), listed last and marked "[TC]" in 
Table II. The left cheliped is normal, as are the right merus 
and carpus. However, the inferior margin and the lower part 
of the outer surface of the right (major) propodus were ap- 
parently slashed or constricted distally, and although the 
wound healed completely, a depressed scar remains, the distal 
proportions of the chela are strongly altered, and ornamenta- 
tion on the outer surface of the propodus is confined to small, 
sharp, scattered projections. Also, extension is restricted be- 
tween carpus and propodus due to incompletely developed 
hinges. 

Discussion 

Reversed asymmetry of chelipeds has been described for 
some ocypodid brachyurans, alpheid shrimp, and nephropid 



72 Proceedings of the Biological Society of Washington 

macrurans. Details of reversal vary from group to group, but 
perhaps the strongest contrast to the reversal in Calappa is 
Herrick's (1907) account of the approximately equal frequency 
of complete and perfect reversal of asymmetry on chelae 
of the American lobster, Homarus americanus H. Milne-Ed- 
wards, 1837. Herrick (1907, p. 275) indicated that handedness 
on these animals is a genetically controlled, inherently vari- 
able but not random character, and stated that "this condition 
is probably one of direct inheritance, all members of a brood 
being either right-handed or left-handed," the position of the 
major ( "crushing" or "club" ) and minor ( "toothed" or "quick" ) 
chelae being "predetermined in the egg." 

Whereas reversal may be frequent, complete, and perfect on 
lobsters, it is infrequent and imperfect on specimens of Ca- 
lappa. Because features unlike those normally observed on un- 
reversed chelipeds occur on the individual elements of reversed 
chelipeds, complete reversal in these crabs is attributed here 
to simultaneous and equal or near-equal regeneration of both 
chelipeds following autotomy. In support of this interpreta- 
tion, it should be noted that these appendages are almost equal 
in size but are generally slightly smaller than their normal, un- 
reversed counterparts, and their individual elements are more 
or less correctly proportioned relative to one another but, again, 
are somewhat smaller than their unreversed counterparts. 

Thus, when both chelipeds are lost on specimens of Calappa, 
their asymmetry may be reversed completely during subse- 
quent regeneration, and occurrence of this phenomenon is not 
confined to either sex; if examined only superficially, the left 
cheliped usually appears almost perfectly formed, but the right 
is generally malformed in several respects, most noticeably in 
size and in the characteristics of the wing of the merus and 
the proportions and ornamentation of the propodus. Truly 
near-perfect reversal of the asymmetry of chelae was observed 
only on C. hepatica (USNM 93819), but even this specimen 
lacks some, albeit minor, features on its chelipeds which are 
diagnostic of a normal, unreversed condition. 

It may be significant that when only one cheliped is lost, 
the attributes of the replacement chela are apparently never 



Asymmetry of Calappa chelae 73 

those associated with the original major chelipeds. Regardless 
of which cheHped is autotomized, the plan of the regenerated 
appendage seems to be the simpler of the two possible alter- 
natives, but one should probably not attach phylogenetic sig- 
nificance to this observation, especially since at present the 
genetic and/or biochemical mechanisms whereby the charac- 
teristics of replacement appendages are determined are so 
poorly understood. 

Regarding general ramifications of this study, it is suggested 
that autotomy and subsequent regeneration of chelipeds among 
Recent species of Calappa is comparatively uncommon, partly 
because their digging or burrowing habit affords the animals 
considerable initial protection, but primarily because the pos- 
terolateral clypeiform expansions and the characteristic pos- 
ture with which the chelipeds are carried, closely appressed 
to the frontal and anterolateral margins of the carapace, prob- 
ably provide the most effective protection to these append- 
ages which is to be seen among all brachyurans. This is not 
to imply that the animals are rendered impregnable by their 
tanklike attributes, as three specimens of C. flammea (USNM 
5233 ) , taken in 1883 from fish stomachs off Pensacola, Florida, 
demonstrate conclusively that species of Calappa are preyed 
upon by marine vertebrates. However, because the chelipeds 
are needed to secure food and, at the same time, form an in- 
dispensable part of an elaborate respiratory apparatus neces- 
sarily associated with the animals' normally buried or partially 
buried existence (see Garstang, 1897), it is likely that, follow- 
ing loss of these appendages, few individuals survive long 
enough to regenerate new chelipeds. 

Because reversal is apparently associated only with regenera- 
tion, it appears that under all normal conditions the major 
chela in Calappa occurs only on the right cheliped and that this 
character is probably ancient, well established, and strongly 
maintained genetically. As reversal per se is presumably neither 
advantageous nor disadvantageous to an individual, its sur- 
vival value is probably negligible so long as one cheliped pos- 
sesses the unusual characteristics of a major chela in perfect 
working order. The two specimens whose major chelae were 



74 Proceedings of the Biological Society of Washington 

not fully functional (C. flammea, USNM 71113 and C. he- 
patica, USNM 65418) probably experienced difficulty in feed- 
ing using the techniques described by Shoup (1968). How- 
ever, the two individuals which exhibit partial reversal and 
lack major chelae on both chelipeds (the male C. sulcata, 
USNM 103450 and C. hepatic^ USNM 99134) were certainly 
prevented from feeding in their characteristic manner on their 
preferred diet of gastropods and to have survived, they must 
have altered their feeding habits considerably. 

This study was concerned initially only with species of Ca- 
lappa, but chelae reversed according to the previously noted 
general rules occur also in species of other calappid genera. Re- 
versal is complete and nearly perfect on a male specimen of 
Cycloes hairdii Stimpson, 1860, collected off Sao Luis, Brazil, 
in 1963 (USNM 123339); both chelipeds are similar in size, 
shape, proportions, and ornamentation, but where only the 
minor (left) propodus customarily bears a stout supradactylar 
projection in this species, strong projections are present on both 
major (left) and minor (right) propodi of this individual. In 
addition, the entire right cheliped has been regenerated on a 
large male specimen of Mursia gaudichaudii (H. Milne-Ed- 
wards, 1837), collected in 1966 northwest of Valparaiso, Chile 
(USNM 123610), and the right (minor) propodus is conspicu- 
ously small and atypical in both proportions and ornamenta- 
tion. Finally, a single, isolated right chela (USNM 77170), 
clearly assignable to Acanthocarpus alexandri Stimpson, 1871, 
and collected in 1930 south of the Tortugas, Florida, was re- 
generated grotesquely as a minor chela. Thus, complete and 
partial reversals evidently occur throughout the Calappinae, 
but judging from the small percentage of specimens of Calappa 
which exhibited reversed chelae, these phenomena are prob- 
ably not widespread within either a population or any given 
species. 

Reports of reversal in Calappa are rare in the literature and 
where they exist, they are brief statements without detailed 
descriptions or discussion. Holthuis (1958: p. 172) cited a 
probable case of partial reversal on a male specimen of the 
Caribbean species C. cinerea Holthuis, 1958, from St. Martin, 



Asymmetry of Calappa chelae 75 

Netherlands Antilles. The right chela he described as "very 
small and of abnormal shape, somewhat resembling the small 
[left] chela," concluding, "It is evidently in process of regen- 
eration." 

The only other report of reversal known to the writer is that 
by Barnard (1950: p. 347), who mentioned having seen "two 
'sinistral' specimens in 20 specimens" of C. hepatica spinosis- 
sima H. Milne-Edwards, 1837. Barnard did not specify whether 
reversal on these specimens is complete or partial, although by 
noting this occurrence in the same short footnote with com- 
ments concerning the seemingly perfect reversal on Miers' 
(1886) plates of the "Challenger" brachyurans, he implied that 
reversal on these specimens is complete. If this was indeed his 
intention, the 10 percent reversal reported for C. hepatica 
spinosissima represents a significantly greater frequency than 
that observed on specimens in the Smithsonian collections, 
in which slightly less than 1 percent of all specimens are 
thus affected. Of the 172 specimens of C. hepatica exam- 
ined, only two, or a little over 1 percent, have completely re- 
versed chelae, and neither of the two specimens of C. hepatica 
spinosissima exhibits reversal. Therefore, either the specimens 
studied by the writer do not accurately reflect the frequency 
of the phenomenon in nature, or Barnard's samples were atypi- 
cal, or complete reversal is unusually common within this sub- 
species, at least from this geographic area. It is impossible to 
determine which, if any, of these alternatives is correct, but 
because of its diversity and large numbers of individuals, the 
material in the Smithsonian collections is considered fairly 
representative of the genus. 

Regarding the apparent reversal in Miers' (1886) illustra- 
tions of the chelae of C. flammea (pi. 23, fig. lb) and C. de- 
pressa Miers, 1886 (pi. 23, fig. 2b), Barnard (1950: p. 347) 
observed that these figures "may be mirror-pictures due to re- 
versal in the lithographing process." That Barnard was proba- 
bly quite correct is indicated by the total lack of comment con- 
cerning reversal in Miers' text; had any of the "Challenger" 
specimens exhibited such an unusual phenomenon, it is ex- 
tremely unlikely that Miers would have failed to mention it. 



76 Proceedings of the Biological Society of Washington 

Indeed, Miers' illustrations of the chelae of Paracyclois milne- 
edwardsii Miers, 1886 (pi. 24, fig. Ic), and Mursia curtispina 
Miers, 1886 (pi. 24, fig. 2c), likewise representatives of the 
Calappinae, also show apparent reversal, but again no rele- 
vant comments appear in the text. 

One of the lithographs of Calappa granulata (Linne, 1758), 
executed by Roux (1828), shows chelae apparently reversed 
(pi. 2, fig. 1), but another in the same work shows them in 
unreversed position (pi. 16, figs. 1, 3-4). The excellent detail 
in the former indicates that in size and proportions, at least, 
these elements are quite normal, but, as in Miers ( 1886 ) , there 
is no mention of reversal or variability in handedness in the 
text, and this is probably another example of error attributable 
to the lithographing process. 

Summary and Conclusions 

In the Calappinae, major chelae bearing specialized struc- 
tures normally occur on the right chelipeds and minor chelae, 
lacking these structures, on the left. To investigate the exis- 
tence, frequency, and significance of reversed asymmetry of 
chelae in Recent species of Calappa and to assess the degree of 
variability and malformation of their chelipeds, the writer ex- 
amined 809 identified specimens in the collections of the Smith- 
sonian Institution and found 15 with abnormal chelipeds. 

Complete reversal of asymmetry was observed on both cheli- 
peds of seven specimens, and in these cases, reversal is at- 
tributed to simultaneous and approximately equal regeneration 
of these appendages following autotomy rather than to any 
inherent genetic causes. Features which usually occur only on 
completely reversed chelipeds and differ from those on normal, 
unreversed chelipeds include the following: (1) on both cheli- 
peds: smaller than normal size, relative to size of carapace and 
other appendages; (2) on left (major) chelipeds: (a) reten- 
tion of all or part of supradactylar projection on propodus, and 
(b) virtual absence of elevated tooth on superior proximal 
margin of dactylus; and (3) on right (minor) chelipeds: (a) 
smaller size of entire chelipeds and individual elements, rela- 
tive to left counterparts and to minor cheliped on unreversed 
specimens, (b) reduced number of teeth on wing of merus, 



Asymmetry of Calappa chelae 77 

strength and delineation of individual teeth being generally 
weak and overall appearance of wing aberrant, (c) atypical 
shape and proportions of propodus, and (d) paucity of char- 
acteristic ornamentation on outer surfaces of all elements, espe- 
cially on propodus, where occasional random tubercles and 
patches of granules occur on otherwise smooth surface. These 
characters apply well to all seven reversed specimens except 
in the ornamentation and proportions of the right (minor) 
propodi on two specimens of C. hepatica which exhibit al- 
most perfect reversal. 

When only one cheliped is regenerated, the plan of the re- 
placement chela is apparently always that of the minor, more 
generalized chela, regardless of whether it is produced on the 
left or right appendage; if on the latter, the condition consti- 
tutes partially reversed asymmetry. Features exhibited on seven 
specimens, characteristic of regeneration of only a single 
cheliped, include: (1) quite normal size, shape, proportions, 
and ornamentation of unregenerated cheliped and its elements, 
in contrast to markedly smaller size and deviations in shape, 
proportions, and ornamentation of regenerated limb and its 
parts, (2) reduced number and atypical ornamentation of 
teeth on wing of regenerated merus, (3) strongly aberrant 
shape and proportions of regenerated propodus, (4) generally 
smooth outer surface of propodus, with few, knobby, usually 
randomly distributed tubercles, and (5) unusual characteris- 
tics of superior crest of propodus, either dentate with an ex- 
cessive number of teeth or rounded with no teeth at all. 

The one injured specimen which the writer encountered 
suggests that when an element of a cheHped is damaged, the 
wound may affect subsequent growth and the involved part 
may become conspicuously atypical and, depending on the 
extent of the injury, even grotesque. However, the abnormaHty 
is generally confined to that particular element and, therefore, 
the condition is easily recognized. 

With the demonstration of complete and partial reversal of 
asymmetry of chelae in species of the Calappinae, it becomes 
obvious that ordinarily no taxonomic significance can be at- 
tached to the attributes of reversed or injured chelae on speci- 



78 Proceedings of the Biological Society of Washington 

mens otherwise clearly assignable to this subfamily. Because 
the fossil propodus mentioned earlier probably represents a 
case of complete and nearly perfect reversal, it may represent 
an exception to the generalization just stated, and its attributes 
may be taxonomically significant at the specific level. Conse- 
quently, the type-species of the fossil genus recognized by 
Ross, Lewis, and Scolaro ( 1964) should probably be reassigned 
to the genus Calappa, with which it has its closest affinities. 

Acknowledgments 

Financial support for the initial part of this study was re- 
ceived from U.S. PubHc Health Service grant 5-TOl ES 00027- 
04, administered by the Tulane University Program in Environ- 
mental Biology. A predoctoral internship from the Smithsonian 
Research Foundation enabled the writer to return to the Mu- 
seum of Natural History where he completed the study. For 
both of these awards, the writer is extremely grateful. In addi- 
tion, deep appreciation is extended to Dr. R. B. Manning and 
Dr. T. E. Bowman of the Smithsonian Institution, who gener- 
ously authorized access to the collections and provided work 
space at the museum during the summer of 1967. Finally, he 
is pleased to acknowledge with deep thanks the counsel of Dr. 
F. A. Chace, Jr., and Messrs. H. B. Roberts and E. S. David- 
son, whose suggestions assisted the writer in reorganizing and 
clarifying parts of the original manuscript. 

Literature Cited 

Alcock, Alfred. 1896. Materials for a carcinological fauna of India. 
No. 2. The Brachyura Oxystoma. Jour. Asiatic Soc. Bengal, 
65 (2) (2): 134-296, pis. 6-8. 

Barnard, K. H. 1950. Descriptive catalogue of South African dec- 
apod Crustacea (crabs and shrimps). Ann. South African 
Mus., 38: 1-837, 154 figs. 

Garstang, Walter. 1897. On some modifications of structure sub- 
servient to respiration in decapod Crustacea which burrow in 
sand; with some remarks on the utility of specific characters 
in the genus Calappa, and the description of a new species 
of Albunea. Quart. Jour. Micro. Sci. (N.S.), 40 (2): 211- 
232, pis. 12-14. 

Herrick, F. H. 1907. Symmetry in big claws of the lobster. Science, 
25: 275-277. 

HoLTHUis, L. B. 1958. West Indian crabs of the genus Calappa with 



Asymmetry of Calappa chelae 79 

a description of three new species. Stud. Fauna Curagao 

Caribbean Isls., 8 (34): 146-186, figs. 28-54, tbl. 14. 
Lewis, J. E. 1969. Aparnocondylus Ross, Lewis, and Scolaro, 1964 

(Decapoda: Oxystomata ) : a reappraisal. Jour. Paleontology 

[in press]. 
MiERS, E. J. 1886. Report on the Brachyura collected by H.M.S. 

"Challenger" during the years 1873-1876. Rept. Sci. Res. 

Challenger (Zool.), 17 (49): L + 1-362, 29 pis. 
Rathbun, Mary Jane. 1937. The oxystomatous and allied crabs of 

America. U.S. Natl. Mus. Bull. 166: vi + 1-278, 86 pis., 47 

figs., 87 tbls. 
Ross, Arnold, J. E. Lewis, and R. J. Scolaro. 1964. New Eocene 

decapods from Florida. Quart. Jour. Florida Acad. Sci., 27 

(3): 187-196, 2 figs. 
Roux, PoLYDORE. 1828-1830. Crustaces de la Mediterranee et de 

son httoral decrits et lithographies par Polydore Roux. Paris 

& Marseille: 176 pp. [n.n.], 45 pis. 
Shoup, J. B. 1968. Shell opening by crabs of the genus Calappa. Sci- 
ence, 160 (3830): 887-888, 3 figs. 



80 Proceedings of the Biological Society of Washington 



4 



73 



82, pp. 81-92 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



RHAPTOTHYREUS TYPICUS N.G, N.SP., AN ABYSSAL 

MARINE NEMATODE REPRESENTING A NEW 
FAMILY OF UNCERTAIN TAXONOMIC POSITION 

By W. D. Hope and D. G. Murphy* 
Smithsonian Institution, Washington, D. C. 

Several unusual male and juvenile specimens of a new spe- 
cies of abyssal, marine nematode have been collected near the 
Pacific coast of Colombia and Chile, off the southwest coast of 
South Africa, and off the east coast of Florida by the Lamont 
Geophysical Laboratory; and on the Atlantic slope near the 
coast of New Jersey, and at several stations near Bermuda by 
the Woods Hole Oceanographic Institute. 

These nematodes are striking in that they possess an excep- 
tionally large, oval amphid with a median longitudinal rib, 
a vestigial stoma and esophagus, no apparent gut, and a single 
spiculum. Notwithstanding their wide geographic separation, 
the specimens from all of these stations have no morphological 
differences judged to be interspecific. 

Females may also be present in the same collections but, if 
so, they are not recognizable because of pronounced sexual 
dimorphism. Examination of the juveniles revealed no clues 
to the identity of females. 

The greatest abundance of specimens from a single locality 
was found off the coast of New Jersey, and it is upon this popu- 
lation that the following description is based. 

All but two specimens were prepared as permanent whole- 
mounts in glycerine; the remaining two were embedded in 
polyethylene glycol, sectioned at 5 fx, and stained with hema- 
toxylin by the method of Craig and Wilson ( 1937 ) . 



'= Visiting Research Associate. 

5— Prog. Biol. Soc. Wash., Vol. 82, 1969 (81) 






82 Proceedings of the Biological Society of Washington 

RHAPTOTHYREIDAE new family 

Diagnosis: Rather large nematodes. Cuticle smooth but with very fine 
transverse striations observed in some regions. Somatic papillae very 
small and sparse. Amphid oval, 2 to 3 head diameters long, with median 
longitudinal rib and transverse rows of punctations. Stoma and esophagus 
vestigial, gut apparently absent. Spiculum single and gubernaculum 
absent. Taxonomic position uncertain. 

Rhaptothyreus new genus 

Diagnosis: Same as that of the family. The name Rhaptothyreus is 
descriptive of the amphid and is derived from the Greek rhaptos, meaning 
stitched or sewn, and thyreos, meaning a large, oblong, door-shaped 
shield. 

Rhaptothyreus typicus new species 

Specimens: 18 males; 2 juveniles. 

Measurements of males: 

Holotype: L = 9.80 mm; a = 87.4; b = 33.0; c = 133.0. 

Paratypes: L = 6.99-12.80 (8.79 ± 1.26) mm. 

a = 67.2-98.8 (78.9 ±6.9). 

b = 26.5-44.4 (32.7 ±4.0). 

c = 100.6-152.0 (122.2 ± 14.1). 

Description: Body slender and gradually tapered anteriorly; posteri- 
orly tapered, curving mostly on dorsal side, from near level of cloaca 



-> 

Fig. 1. Rhaptothyreus typicus n. sp. Wholemount of paratype 
showing the general features of the body including the junction between 
the posterior end of the zone containing basophilic rods and the anterior 
end of the solid cylinder of granular cells (br/sp). X 52. 

Fig. 2. Rhaptothyreus typicus n. sp. A portion of an amphid of the 
holotype depicting the transverse rows of punctations and the longitu- 
dinal rib (Ir). X 1,440. 

Fig. 3. Rhaptothyreus typicus n. sp. Transverse section of a paratype 
cut at the level of the amphids showing the external layer of cuticle on 
the amphid (m) and its longitudinal rib (Ir). X 1,440. 

Fig. 4. Rhaptothyreus typicus n. sp. Transverse section of a para- 
type showing the basophilic rods (br). Note the apparent absence of 
intestinal cells. X 1,440. 

Fig. 5. Rhaptothyreus typicus n. sp. Transverse sections of a para- 
type cut at a level slightly posterior to the junction between the region 
containing the basophilic rods and the cylinder of cells. The body cavity 
is largely occupied by the cells of the cylinder (sp), but a small area 
is occupied by a dense mass of rods (br). Nuclei of the cells constituting 
the cylinder (n sp). X 1,440. 



New marine nematode 83 







br/sp 







84 Proceedings of the Biological Society of Washington 



© 



Ir 





TSlew marine nematode 



85 




86 Proceedings of the Biological Society of Washington 

(Fig, 1). Head diameter at level of cephalic setae 33.0 /a-42.0 /a (37.3 /j- 
+ 2.3 /J,). Body diameter at posterior margin of amphid 60.5 /i-76.0 /x 
(67.7 fji. ± 5.0 fi) ; at base of esophagus 76.5 ^-102.1 ^ (86.6 /. ± 6.1 /x); 
at mid-body length 90.5 /i-133.7 /x (111.3 fi ± 11.1 /x); and at level 
of cloaca 57.5 fi-80.0 fi (67.4 /x ± 6.5 fi). Cuticle without longi- 
tudinal striations, but with extremely fine transverse striae, most evident 
on neck and tail of male and produced by fine transverse ridges and 
furrows on inner surface of cuticle; periodicity of striae 1 yCi-2 fi (Figs. 7 
and 9). 

Twelve cephalic setae, each slightly longer than wide (Figs. 7 and 8 ) . 
Cervical setae also very short and arranged in eight longitudinal rows at 
level of amphid; one row at dorsal and one at ventral margins of each 
amphid and two subdorsal and two subventral rows (Fig. 7). Somatic 
setae equally short, but sparse and not in obvious rows. 

Distance from anterior extremity of head to anterior margin of amphid 
19.0 /x-30.5 fi (25.3 /a ± 3.0 /j.). Amphid a large, oval intracuticular cavity 
83.0 /x-102.0 /x (93.0 fi ± 6.0 fi) long, 31.0 ^-40.0 fi (34.9 /. ± 2.8 /.) 
wide (Fig. 7). External wall of cavity a thin, external layer of cuticle 
bearing closely spaced, transverse rows of punctations and a longitudinal, 
tubular rib on medial surface of membrane ( Figs. 2, 3, and 7 ) . Lumen of 
rib partially divided by discontinuous, longitudinal keel (Figs. 2 and 3). 
Posterior approximately Ys of tube with well-developed, continuous keel 
and two distinct lumina. At posterior end of amphid, each lumen pro- 
ceeds medially into hypodermal tissue. 

Head rounded, without lips or microlabia* (Figs. 7 and 8). Vestige of 
stoma shifted slightly dorsad (Figs. 7 and 8). Neck laterally compressed 
in region of amphid (Figs. 3 and 8). Nerve ring 152.0 /i-193.7 /m (171.6 fi 
± 11.0 fi) from anterior extremity of head. Ventral gland absent. Esopha- 
gus not well-formed, but represented by diffuse, amorphic tissue devoid 
of a lumen ( Fig. 3 ) ; gut apparently absent. Body cavity largely occupied 
by dense aggregations of basophilic, rod-shaped particles approximately 



^ See Hope, 1967, for definition. 



Fig. 6, Rhaptothijreus typicus n. sp. Transverse section of a male 
paratype cut at the level of the spiculum (s). X 1,875. 

Fig. 7. Rhaptothyreus typicus n, sp. Lateral view of the head of 
the holotype. 

Fig. 8. Rhaptothyreus typicus n. sp. Face view of a paratype. 

Fig, 9, Rhaptothyreus typicus n. sp. Lateral view of the tail of 
the holotype. 

Fig. 10. Rhaptothyreus typicus n. sp. Lateral view of the tail of a 
juvenile paratype. 



New marine nematode 



87 




Proceedings of the Biological Society of Washington 




iSlew marine nematode 




90 Proceedings of the Biological Society of Washington 

1 fi long (Fig. 4). Body cavity occupied posteriorly by solid, cylindrical 
mass of large cells with very granular cytoplasm (Figs. 5, 9 and 10). 
Posterior end of cylinder continuous with tube ventral to spiculum; tube 
comprised of small cells and tapered toward posterior end; possibly 
continuous with proctodaeum (Figs. 9 and 10). 

Single spiculum present (Fig. 6), distally arched ventrad and 72.6 /j.- 
147.3 fji (120.4 fi ± 16.0 fx) long. Gubernaculum and copulatory supple- 
ments absent (Fig. 9). 

Cuticle thickened near mid-tail forming narrow, ventral, transverse 
ridge, each lateral end of ridge frequently bearing single minute pore. 
Cuticle at terminus of tail thickened, cap-like (Fig. 9), Caudal glands 
and spinneret absent. Tail length 61.1 ^-84.2 /x (72.0 fi ± 5.9 /x). 

Fourth stage juvenile males closely resembling adult but stoma more 
evident than in male (apparently a narrow tube). Amphid and spiculum 
less cuticularized. Tail bluntly conical, devoid of ventral ridge and 
terminal cap, but inner surface of cuticle on ventral side of tail with 
depression where transverse ridge of adult male tail will form (Fig. 10). 
Contents of body cavity resembling those of adult. Other juvenile stages 
and females not known, 

Holotype: United States National Museum Number 39215. 

Paratijpes: United States National Museum Number 39216-39235. 

Type locality: East of the New Jersey Coast (39° 46.5' N; 70° 
43.3' W) at depths of 1,330 to 1,470 meters. 

Distribution: Specimens, at present judged to be of the same species, 
occurred at the locations given in the table below. 

Discussion: The large size and unusual structure of the amphid, vestig- 
ial nature of the stoma, apparent absence of a functional esophagus and 
gut and the presence of a single spiculum in the male of the above 
described species leave little doubt that these specimens represent not 
only a new genus and species, but a new family as well. On the other 
hand, the uniqueness of these features leave some question as to where 
this family should be placed in the present classification of marine 
nematodes. 

The structure of the amphid of the adult male would suggest that it 
belongs to the class Enoplida, since it is a pouch-like intracuticular 
cavity. From the amphids found in members of the families Leptosoma- 
tidae, Enoplidae and Oncholaimidae it differs in not having a transverse, 
external, slit-like opening at its anterior end. The narrow, oval shape of 
the amphid superficially resembles that found in Halalaimus, but the 
significance of this similarity is difficult to judge since it is not known 
whether the amphid of Halalaimus is an open or a closed cavity. 

The smooth, but finely striated cuticle, and the arrangement and 
number of cephalic setae are also compatible with the structure of the 
cuticle and arrangement of cephalic setae in Enoplida, however, these 
characters do not exclude this family from other major taxa of marine 
nematodes. 



New marine nematode 



91 



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92 Proceedings of the Biological Society of Washington 

The presence of a single spiculum, in itself, does not provide a clue 
to the taxonomic position of this species, since this condition is extremely 
unusual among free-Hving nematodes. 

There are no organs in the body cavity of our specimens readily recog- 
nizable as gut or gonad. That portion of the body cavity usually occupied 
by these organs contains two contiguous zones. The anterior and longer 
one is an aggregate of basophilic rods resembling bacteria, If these are 
bacteria, they possibly are symbionts since they occur in all the speci- 
mens at hand, representing a wide geographic area, and they are within 
the body cavity rather than in the lumen of the gut, suggesting the possi- 
bility of a highly evolved biological relationship. 

The posterior zone is comprised of a compact cylinder of cells, prob- 
ably sperm, having continuity with a duct or vas deferens which opens 
to the exterior of the body near the distal end of the retracted spiculum. 

The juvenile males differ from the adult males in having a less cutic- 
ularized spiculum. Amphids resembling those of the adults are present 
in the juveniles, but they also are less cuticularized and appear to be 
located in the newly developing inner layer of cuticle. Structures that 
could be interpreted as amphids in the outer layer of the cuticle formed 
by the preceding juvenile stage were not seen. Perhaps the amphids of the 
earlier stages are small pores and difficult to resolve. 

The change that takes place in the shape of the tail, and the possibility 
of a change in the structure of the amphid during development, suggest 
that younger juveniles may possess certain morphological features that 
would provide a clue to the phylogenetic relationships of this species. 
Additional clues of relationships may be found in the structure of the 
females which, if present in the same collections as the males, must be 
strikingly different due to sexual dimorphism, as in the Symplocosto- 
matinae. 

Literature Cited 

Craig, R. and C. Wilson. 1937. The use of buffered solutions in 
staining: theory and practice. Stain Technol. 12: 99-109. 

Hope, W. D. 1967. Free-living marine nematodes of the genera Pseu- 
docella Filipjev, 1927, Thoracostoma Marion, 1870, and 
Deontostoma Filipjev, 1916 (Nematoda: Leptosomatidae ) 
from the west coast of North America. Trans. Amer. Microsc. 
Soc. 86 (3): 307-334. 



73 



QH 




1 
B4X 


1. 82, pp. 93-112 29 May, 1969 


NH 


PROCEEDINGS 




OF THE 




BIOLOGICAL SOCIETY OF WASHINGTON 



COLOMBIAN FRESHWATER CRAB NOTES 

By Waldo L. ScHMm 
Smithsonian Institution, Washington, D. C. 

These notes are a rather belated and unorthodox accounting 
of a number of very interesting freshwater crabs sent to the 
U. S. National Museum for identification in 1940-41 by Brother 
Niceforo Maria, Director of the Museum of the Instituto de 
LaSalle, Bogota, Colombia, who had collected them from as 
far south at Puerto Asis, Int. de Putumayo, not far from the 
Ecuadorian border, to Cucuta and vicinity, Norte de Santander, 
close to the Venezuelan border. The specimens in the Niceforo 
collection are commented upon below under their current 
species name headings. Among them were found four new 
species and a new subspecies, of which all but the one de- 
scribed in this paper were published upon by Dr. Gerhard 
Pretzmann (noted below). A number of new records of occur- 
rence for these Colombian freshwater crabs are here estab- 
lished. 

Concluding these notes are some remarks on variation in the 
carapace armature of Valdivia (Valdivia) piriformis Pretz- 
mann, and, though not represented in the Niceforo material, 
of Trichodactylus (Dilocarcinus) pictus (A. Milne-Edwards, 
1853) (<cHolthuisia pictus). 

The figures of the gonopods and carapaces of most of the 
species listed or discussed are, in the case of the gonopods, 
unless otherwise stated in the figure legends, three times nat- 
ural size, and in the case of the carapaces close to natural size. 

Family Trichodactylidae 

Valdivia (Rotundovaldivia) niceforoi (Schmitt and Pretzmann, 1968) 

Figure 1, a-d 

The first of the new species is a trichodactylid which, in the char- 
acter of its gonopods, is unhke any member of the family previously 

6— Prog. Biol. Soc. Wash., Vol. 82, 




94 Proceedings of the Biological Society of Washington 




Fig. 1. Valdivia (Rotundovaldivia) niceforoi (Niceforo No. 4; USNM 
112117). a. right gonopod, posterior view. b. same, reversed, anterior 
view. c. second right gonopod. d. carapace. 



known. Dr. Gerhard Pretzmann, of the Naturhistorisches Museum, 
Vienna, at the time engaged in reviewing the taxonomy of American 
freshwater crabs, visited Washington in October 1964 for the purpose 
of examining pertinent material in the National Museum. 

He became much interested in this new species, as well as in the 
other identifications that I had made of the Niceforo crabs, and wished 
particularly to include mention of the new species in a forthcoming ac- 
count of his recent studies. Because of procrastination on my part and 
preoccupation with more pressing commitments, I acceded to his pub- 
lishing it with the understanding that it be named for Brother Niceforo. 

Dr. Pretzmann subsequently published a diagnosis and brief description, 
in the authorship of which he included me: "Trichodactylus (Valdivia) 
niceforei Waldo L. Schmitt and Gerhard Pretzmann" [1968b, p. 61]. 
The specific name, however, should be spelled niceforoi. 

Though I would have wished it otherwise. Dr. Pretzmann selected 



Colombian freshwater crab notes 95 

for the type a specimen that Brother Niceforo had sent to the Paris 
Museum at an earlier date: "Typus: $ ? 32.1 mm Cpxlg. Fundort: 
Pamplona. M. Nicefore coll. 1936." Whether that sex-questioned speci- 
men was a damaged male or one from which the gonopods were missing 
was not made clear. 

Not quite two months later Dr. Pretzmann removed the species to his 
newly created subgenus Rotundovaldivia of the genus Valdivia [1968c, p. 
73]. 

Inasmuch as the Niceforo specimen in the U. S. National Museum 
that had alerted me to this new species is no more than a shade larger 
than the one that Dr. Pretzmann designated the type of the species, 
34 as compared to 32.1 mm in median length, I am moved to contribute 
here a somewhat more detailed description of the species based on the 
National Museum specimen, together with some remarks on three other 
males and two females collected by Brother Niceforo while resident for 
a time in the Colegio del Corazon, Ciicuta, Norte de Santander. 

Description of my specimen of V. (R.) niceforoi, 34 mm in median 
length of carapace from Cucuta, Norte de Santander (Niceforo No. 4, 
USNM 112117): 

Carapace longitudinally very convex, more oval than suborbicular, 
about three-fourths as long on median line as greatest width across tips 
of posterior pair of lateral spines. Front bilobed, margins of lobes mi- 
nutely roughened with fine granules, as is general surface of the cara- 
pace; median emargination of front about one-third as deep as the orbit 
in dorsal view measured back from a Hne joining the anterior margins of 
the unarmed frontal lobes; at its middle the frontal emargination 
is armed with a pair of sharp, narrowly triangular teeth or spines, on 
either side of which there is a small tooth or tuberculiform denticle; 
viewed from below these denticles are more conspicuous, suggestive of 
a second incipient pair of frontal spines. The adjacent margins of the 
frontal lobes, furthermore, tend toward being tuberculated, due, appar- 
ently, to an enlargement of the granulations of the carapace as they 
reach the frontal margin. 

Outer angle of orbit armed with a sharply triangular tooth, about as 
far removed from the first or anterior of the seven anterolateral spines as 
the last is from the one that just precedes it; the first anterolateral tooth 
is about as far removed from the second as the fifth is from the sixth; 
interspaces between third, fourth, and fifth spines subequal. Ventral 
margin of the orbit armed with five somewhat medially directed blunt 
spines or elongate denticles, of which the innermost is the largest (tip 
of the corresponding one on the left side is broken off); thickened 
margins of the median projection of the epistome at apex of projection 
(in dorsal view) appear to form either side of the median line a low 
swelling; anterolateral angle of buccal frame armed with six somewhat 
elongate-tuberculiform denticles on the right side, five on the left. 

The chelae are markedly unequal in the unique type. The right one is 
vastly the larger, the perpendicular (greatest) height being more than 



SMVUISONIAN wi\Y29TO 
\KSini)Tinr 



96 Proceedings of the Biological Society of Washington 

twice that of the smaller left chela; in the major chela the length of 
sinus or gape between the fingers is about four-fifths of the distance back 
to midpoint of the posterior margin of the palm proper; in the smaller 
hand the gape is a little greater than the corresponding length of the 
palm; upper margin or surface of either palm flattened, bluntly angu- 
lated where it passes over into inner and outer faces; ventrally the palmar 
margin is evenly rounded off. Carpus of either cheliped carries but a 
single stout spine on the inner margin, arising about one-third the length 
of the carpus from the anterior end; it is a little more slender and sharper 
on the minor carpus; anterior margin of upper surface of merus armed 
in the major cheliped only with a low spine, or tubercle with acute tip 
at about one-third of the length of the margin anterior to the suture 
separating this joint from the ischium, outer margin of merus with a simi- 
lar spine about one-fourth the length of the joint proper from the ante- 
rior end; merus of minor cheliped armed like that of major one, but the 
spines are relatively longer, more slender, and sharper. 

Dactylus of ambulatory legs elongate, triangular in lateral view, bluntly 
tricarinate; dorsal carinae more or less obscured by dense fringes or 
brushes of short hairs; inner and outer faces flattened and slightly con- 
cave or excavate; ventral margin flattened and bluntly angulated where 
it passes over into inner and outer faces of the dactyl; lower surface and 
marginal angulations of dactyl hidden by a double fringe of hair several 
times as long as the brushes fringing the upper margin of the dactyl; the 
tips of all the dactyls are rather worn off. 

First abdominal somite almost wholly concealed by the carapace, 
second very narrow, third also narrow but twice as long (on median 
line) as second; third to sixth somites fused together, although all suture 
hues externally are very sharply and distinctly marked, so much so that 
all somites to all appearances are freely articulated with one another; 
free seventh somite triangular, lateral margins concave, posterior extrem- 
ity rounded off; this somite is a little more than half as long as wide 
at base. 

The anterior pair of gonopods are unique among those of trichodactylid 
species that so far I have had the opportunity of examining. Much con- 
torted, their distal two-thirds constitutes a hollow sheath or tube through 
which the slender, almost setiform, second pair of gonopods projects as 
much as 2 mm. The inner and underside of the twisted terminal hooked 
portion of the anterior pair of gonopods is furnished with a darker 
colored, minutely spined or prickly area, which, as viewed when in 
position, reappears above on the distal end of the anterior margin of the 
"hook" of the gonopod. 

Measurements: The median length of the carapace of the specimen 
described above, measured from between the pair of spines in the frontal 
emargination, is 34 mm; the width over the apices of the anterolateral 
spines is 45.2 mm; width between apices of spines arming outer angle 
of orbits 25 mm; the dorsal margin of major palm 19 mm long, the mov- 



Colombian freshwater crab notes 97 

able finger 27 mm; the ventral margin of hand from articulation to tip 
of immovable finger 43 mm. The corresponding measurements of the 
minor chela are 10.0, 18.5, and 28.5 mm. The terminal seventh abdominal 
somite is 6 mm long by 11 mm wide. 

Remarks: The five other specimens (three 3 $ and two $ ? ) of 
this species at hand conform in general appearance and in most particu- 
lars with the specimen described above. 

The largest of the three males is 32 mm long and 41.8 mm wide; the 
frontal margin is armed much as in the above described male; the right 
and left lower orbital margins are armed with 4 and 6 spines respectively; 
the corresponding lateral angles have on the one side 3 spines, with per- 
haps the indication of a 4th spine, and 5 spines on the other. 

The second male is soft (recently moulted) and has a fractured cara- 
pace; it measures 27.4 to 28 mm long by about 35 mm wide; front much 
as in type; 4 and 5 orbital spines; and 5 spines at either angle of buccal 
frame. The third and smallest male is 22.2 mm long by 29.9 mm wide; 
the pair of spines in the median emargination of the front are relatively 
a Httle more separated than in the other specimens; orbital spines 5 
either side; at buccal angles, 4 and 5. 

In general, the female of the species differs but little from the male. 
The female carapace does not taper as much posteriorly, and so in out- 
line is a bit more squarish than that of the male. The armature of the 
median frontal emargination, which in the female tends to depart from 
the male norm of two teeth, is perhaps a variable character, to judge 
from the two females at hand; in the larger one, as noted above, the 
emargination is armed with three teeth, in the smaller specimen with 
one only. 

The chelae of the females are much less developed than those of the 
males. On the other hand, the spines with which the several joints of the 
chelipeds are armed are more prominent, more slender, and very sharp. 
Above the inner articulation of the movable finger of the major chela 
there is a short, conical, acute spine; in the largest of the three male 
specimens there is just an inconspicuous, low, blunt swelHng in this 
position; only the smallest male has a conical, but less slender, spine 
than in the female. The corresponding spine of the minor chela is con- 
ical-tubercuhform in the larger males and in the smallest one more 
slender and more sharply pointed, as is the similarly placed spine in the 
females. 

The larger of the two females is about the size of the described male, 
34.2 mm long by 42.1 mm wide. In the frontal emargination there are 
three sharp spines, a fairly long, more or less centrally placed spine, with 
a slightly smaller one about two-thirds the size of the median spine 
either side of it; the denticle remarked in the large described male 
either side of the spines arming the frontal emargination is of good size 
and more conspicuous than in any of the other specimens at hand; or- 
bital spines are five either side; at buccal angles right and left, 5 and 4. 
The carapace of the smaller female measures 33 by 41.6 mm. In this 



98 Proceedings of the Biological Society of Washington 

specimen the frontal emargination is armed with but a single spine in 
place of the usual median pair; a pair of frontal denticles, as in the other 
specimens, is present. 

Yaldivia (Rotundovaldivia) niceforoi cucutensis Pretzmann 1968 

As remarked above, the more or less roughened margins of the frontal 
emargination of our specimens either side of the aforementioned denticles 
verge on becoming tubercuHform. It is not inconceivable that with 
further development such "tubercles" could become more or less spine- 
hke and so enter into a count of what might be considered a part of the 
spination of the front or its emargination. I am inclined to believe that 
something of this sort may have encouraged Dr. Pretzmann to describe 
as a new subspecies a specimen he had seen in the Basel Museum as 
Yaldivia {Rotundovaldivia) niceforei [sic] cucutensis, "Holotypus: 5 
33.5 mm Cpxlg., Museum Basel, Nr. 763-b. Fundort: Columbien, 
Cucuta" [1968, p. 73]. 

Yaldivia {Yaldivia) serrata White, 1847 
Figure 2, a-e 

In validating the genus Yaldivia White, 1847, of which Yaldivia 
serrata is the type, Pretzmann [1968c, pp. 71, 73] set up two subgenera, 
Yaldivia and Rotundovaldivia, the latter with "Typus subgeneris: T. 
bourgeti." In it, along with several other species, he placed Y. (R.) 
niceforoi, described above. For the subgenus Yaldivia, to which he as- 
signed "y. devillei" and several other known species, he described two 
new subspecies of serrata and two new species: V. (V.^ torresi from 
"Kolumbien, La Regla" and V. (Y.) piriformis (discussed below) from 
"Cucuta." Inasmuch as no illustrations accompany any of the descrip- 
tions of these new forms of Pretzmann's, I shall not undertake to attach 
a subspecific name to three specimens in the Niceforo collection which 
I take to be Y. (V, ) serrata. All three are from the same locality: Puerto 
Asis, Int. de Putumayo," a male (No. 12, 48 X 55.6 mm) and two females 
(No. 13, 42.7 X 49.2 mm; and No. 14, 38 X 45.5 mm). 

Yaldivia (Yaldivia) piriformis Pretzmann, 1968 
Figure 3, a-e, and f-i 

In the Niceforo collection were four male and five female specimens 
which I believed represented Dr. Mary J. Rathbun's Trichodactylus 
venezuelensis, having compared a female of similar size with the cotype 
female in the U. S. National Museum, making allowances for what was as- 
sumed to be variation. No males, so far as I was aware, had at that time 
been linked with certainty to the Rathbun cotypes. Nor was there an 
identified male supposedly of this species available in the Paris Museum, 
where the type was examined for me two years ago by Mr. Edward 
Davidson, of the U.S.N.M. 

During his Washington visit. Dr. Pretzmann, who examined these 
specimens, added to each bottle a note: "Trichodactylus (Yaldivia) 



Colombian freshwater crab notes 



99 




Fig. 2. Valdivia (Valdivia) serrata (Niceforo No. 12). a. right 
gonopod, posterior view. b. same, reversed, anterior view. c. same, medial 
view. d. second right gonopod. e. carapace. 



devillei, Pretzmann, Oct. 1964," without any indication that an unde- 
scribed species might be involved. V. (V.) devillei (A. Milne-Edwards), 
by the way, had been founded on a dried specimen of a female, and, as 
in the case of Miss Rathbun's venezuelensis, Mr. Davidson found no males 
in the Paris Museum that had been identified with this species. 



100 Proceedings of the Biological Society of Washington 








Fig. 3, a-e. Valdivia (Valdivia) piriformis, a. right gonopod (Nice- 
foro No. 24), posterior view. b. same, reversed, anterior view. c. second 
right gonopod. d. carapace of smallest male (Niceforo No. 19). e. cara- 
pace of largest female (Niceforo No. 22). 



On his return to Vieima, however, Dr. Pretzmann described his Val- 
divia (Valdivia) piriformis, "Holotypes: $ 48 mm Cpxlg., Mus. Washing- 
ton (als devillei bezeichnet). Fundort: Cucuta. Nicefore coll." [1968c, 
p. 73]. 

I have no doubt that the cited type is Niceforo's specimen (No. 23) 
from Sardinata, northwest of Cucuta, to which Pretzmann had added a 
devillei label; it is a male a shade over 49 mm long by 51 mm wide; 
two other males ( No. 24 ) from the same locahty measure 32 and 37 mm 
long. Niceforo's collection also includes three females 36, 37, and 40 mm 



Colombian freshwater crab notes 



101 






Fig. 3, r-i. Valdivia (Valdivia) piriformis, f. carapace (Niceforo 
No. 24, smaller specimen), g. carapace (Niceforo No. 24, larger speci- 
men), h. carapace (Niceforo No. 20). i. carapace (Niceforo No. 23). 
As figured these carapaces approximate %o natural size. 



long, and a small male 22 mm long, all four of which (No. 19) were 
collected close to the Venezuelan border north of Ciicuta; a female ( No. 
15) from Rio Caqueta, Int. de Caqueta in the far south of Colombia 
(each of the foregoing specimens measures 2 mm, more or less, wider 
than long); and, lastly, the largest (and widest) of all the specimens 
(No. 22), a female 54.5 mm long by 59.2 mm wide from Puerto Salgar, 
Rio Magdalena, Cundimarca. From the type locahty, Ciicuta, also comes 
a male (No. 20) 44.2 mm long by 45.9 mm wide. 

? Valdivia (Valdivia) devillei (A. Milne-Edwards, 1853) 

In the Niceforo collection is a lone female which I took to be T. (V.) 
peruviarms (A. Milne-Edwards, 1869) (51.8 X 56 mm) to which Dr. 
Pretzmann also attached a devillei label, but whether he now considers it 
another specimen of V. (V.) piriformis must await the pubHcation of his 



102 Proceedings of the Biological Society of Washington 







Fig. 4. Hypolobocera (Neostrengeria) macropa (Niceforo No. 6). 
a. left gonopod, posterior view. b. same, from slightly different angle, 
c. same, reversed, anterior view. d. second right gonopod. e. terminal, or 
apical, funnel of first gonopod from above, f. carapace. 



more complete account of the results of his Washington visit. 

Pretzmann has placed T. (V.) peruvianus, along with T. (V.) vene- 
zuelensis, in his more recently diagnosed new genus Holthuisia, for which 
he selected Dilocarcinus pictus A. Milne-Edwards, 1853, as the geno- 
type [1968c, p. 74]. (The spelUng Holthuisisia in the generic heading 
is probably the result of a typographical error.) 

Family Pseudothelphusidae 
Hypolobocera ( Hypolobocera ) bouvieri ( Rathbun, 1898 ) 

Of this species there were five specimens, a male without specific 
locality data (35.9 X 57.1 mm), and four females; the first of the females 



Colombian freshwater crab notes 103 

(No. 5, measuring 18 mm in median length of carapace X 32 mm wide) 
is from San Gil, south of Bucaramanga; the second (No. 7, 46 X 74.8 
mm) is from Sasaima, northwest of Bogota; the third (No. 11, a badly 
broken specimen whose length could not be determined but which may 
have been between 64 and 65 mm wide) is from Rio Caquita, Int. de 
Caquita; the fourth (No. 18, 45.2 X 73.2 mm) is from Chinacota, 42 
kilometers south of Cucuta. Dr. Pretzmann concurred with these identi- 
fications. 

Hypolobocera (Neostrengeria) macropa (A. Milne-Edwards, 1853) 
Figure 4, a-f 

Of this species there was but a single specimen, fortunately a male 
(No. 6, 31 X 52 mm) from Bogota, unmistakably this species. 

Hypolobocera (Neostrengeria) guenteri (Pretzmann, 1965) 
Not until notice of it appeared in print did I become aware that Dr. 
Pretzmann had described a third new species from among the Niceforo 
specimens, which he named Strengeria (Neostrengeria) guenteri: "Typus: 
$ 25 mm Cpxlg., U. S. National Museum. Locus typicus: Columbien 
160/623, Nikiforov coll." [1965, p. 7]. This specimen (Niceforo No. 8) 
is from Bogota, but, as the type is still in Dr. Pretzmann's possession, I 
cannot now ascertain whether I had essayed an identification of it or 
not. Somewhat later Dr. Pretzmann discovered that his new genus 
Strengeria was but a junior synonym of Hypolobocera [1968a, p. 2]. 
Besides the holotype. Brother Niceforo collected two other specimens 
(No. 9, 26.5 X 44.3 mm and 28.4 X 49.9 mm) at Pennsilvania, Caldas. 

Hypolobocera (Phyllothelphusa) niceforoi new species 
Figure 5, a-e 

Seldom do the descriptions of American freshwater crabs, aside from 
the gonopods, offer the information needed for the precise determination 
of their species. Is it not a fact that species, subgenera, and even genera 
can be distinguished with certainty only on the basis of the nature and 
character of their gonopods? It is most unfortunate that these have yet 
to be adequately classified and so described that they may be keyed out 
and taxonomically recognized or visuahzed from printed (or written) 
text. A promising advance toward the solution of this problem was made 
by Dr. Alfred E. Smalley [1964a] in "A Terminology for the Gonopods 
of the American River Grabs." It is to be hoped that he will continue 
this Hne of investigation, for it has become more necessary than ever in 
view of the regrettably unillustrated reports that Dr. Pretzmann has 
pubHshed over the past several years in his study of various collections 
of South American freshwater crabs in this country and abroad. There 
certainly must be an evolutionary pattern or sequence of some sort in 
which there will be found a number of fundamental gonopod types about 
which all other gonopods can be grouped or arranged in rational hues 
of relationship. 



104 Proceedings of the Biological Society of Washington 




Fig. 5, a-e. Hypolobocera (Phyllothelphusa) niceforoi new species 
(USNM 125133). a. right gonopod, posterior view. b. same, reversed, 
anterior view. c. second right gonopod. d. terminal, or apical, funnel of 
first gonopod from above, e. carapace. 

Fig. 5, f-h. Hypolobocera (Phyllothelphusa) lindigiana (USNM 
20051). f. right gonopod, posterior view. g. second right gonopod. h. 
carapace. 



Some of the gonopod definitions given by Dr. Pretzmann in his publi- 
cations leave one in a quandary as to the placement of an unknown spe- 
cies among the many subgenera and species he named in them. For this 
reason I have, with some hesitation and misgivings, considered a lone 
male crab from San Gil, south of Bucaramanga, the type locality, as 
representative of Pretzmann's [1965] subgenus Phyllothelphusia of his 
genus Strengeria (<iHypolobocera). I have named it in honor of its col- 
lector and donor, and here describe it. 

Description: Carapace moderately convex, finely punctate, with the 
"suture" defining the areolations well marked and broadly troughed; 
front, lightly bilobed, is about one-fourth the greatest width of the cara- 
pace; the extra-orbital angle is quite low and blunt; outer lateral margins, 
though appearing quite smooth, are obscurely crenulate. 

The third maxilHped is almost a facsimile of the same appendage in 
Pseudothelphusa lindigiana as figured by Miss Rathbun, exognath in- 



Colombian freshwater crab notes 



105 







a 

Fig. 6. Pseudothelphusa (Achlidon) agrestis (USNM 19487: fig- 
ures X 5). a. right gonopod, posterior view. b. same, reversed, anterior 
view. c. lateral view of first gonopod. d. second right gonopod. 

eluded; its ischium also is as rectangular in appearance except that the 
anterior margin, if anything, is straighter. 

The chelae, of which the right one is only about two-thirds the size of 
the larger left one, are very similar; the distal half or more of the fixed 
finger of the larger is missing; the movable finger is distally curved 
downward, and, judging from the smaller chela, the curved fingertips 
must have overlapped, leaving no gape between them when closed. The 
movable finger of the larger chela has five fairly large tuberculiform 
teeth, counting from the base; between the pairs of the larger teeth are 
smaller ones grouped in this order and number: 1, 3, 4, 1, while between 
the tip of the finger and the nearest one of the larger tuberculiform teeth 
are five more teeth alternating large and small and all much smaller than 
their counterparts farther back on the finger; toward the fingertip these 
smaller teeth progressively diminish in size. 

The hand and carpus of the cheHpeds are unarmed except for a blunt 
tooth on the carpus well in front of the middle of the margin, which is 
smoothly rounded off; the more or less dorsally turned flattened face of 
the meral joint, which is more or less triangular in cross section, on its 
anterior margin is armed with 64 teeth, all stout and blunt and of fair 
size, except for the diminutive first and last of the row; posterior margin 
of this joint is similarly armed, but with smaller and more numerous 
teeth. 

The ambulatory dactyls, though not very prominently carinated, have 
three rows of distally directed corneous spines above, a median row with 
another row either side of the dorsal margin; a similar but not so regular 
or complete a row of these spines seems to run along each margin of the 
somewhat flattened "lower face" or margin of each dactyl. 

As mentioned above, the gonopods of this species, barring the creation 
of another subgenus for their kind, perhaps best fit into the currently 
monotypic genus Phyllothelphusa, for which Dr. Pretzmann chose Pseu- 
dothelphusa lindigiana Rathbun, 1897, as the "Typus subgeneris." This 



106 Proceedings of the Biological Society of Washington 




Fig. 7. Holthuisia pictus. a. right gonopod (USNM 119881), pos- 
terior view. b. same, reversed, anterior view. c. second right gonopod. 
d. carapace, e, f . carapaces of smallest and largest males ( USNM 125134 ) . 



Colombian freshwater crab notes 107 

subgenus he characterized with the following [1965, p. 8] : "Definition: 
Gonopoden mit einer grossen, blattformigen laterocaudalen Lobe, die 
apical gerichtet ist." If I am correct in this assignment, this characteriza- 
tion of the gonopods would have to be broadened. 

Despite their general conformity to the definition, the gonopods of 
the species here described differ in several particulars from the gonopod 
sketched by Miss Rathbun for Pseudothelphusa Undigiana: They are 
more boldly developed, and the lateral lobes, though somewhat for- 
wardly inclined, are more laterally directed; a slight extension of the 
distal end of a rounded ridge or weal on the "stalk" of the gonopod 
projects a bit over the margin of the lateral lobe, giving the lobe a 
mitten-shaped appearance; the most striking feature of the gonopods, 
however, is the spined margin of the terminal funnel, or "trichter" of 
Pretzmann, as the gonopod is viewed in its normal position on the sternum 
of the crab. Largest and more prominent of the spines arming the distal 
extremity of the gonopod on the uppermost margin of the more or less 
flattened, obhque surface of the apical funnel is a sizable pair with a 
tiny spine or acute denticle between them; beyond this pair are four 
much smaller, sharp, spiniform teeth and on the proximal side a bifurcate 
spine followed down the axis of the gonopod by some smaller irregulari- 
ties. The spination of this funnel margin is better figured than described. 

Measurements: The unique male holotype (USNM 125133) measures 
22.3 mm in median length of carapace by 38 mm wide. The major left 
chela from the tip of the movable finger to the uppermost articulation 
of the hand is about 31 mm long by 11 mm deep, the dactyl or movable 
finger is about 3 mm longer than the 14 mm long dorsal margin of the 
palm; for the smaller left chela the corresponding measurements are 21, 
7, 3, and 12 mm. 

Although I have taken Hypolobocera niceforoi to be representative 
of the subgenus Phyllothelphusa, it is possible that it may fall into the 
subgenus Neostrengeria. Dr. Pretzmann diagnoses the latter [1965, p. 7] 
as follows: "Diagnose: Gonopoden breit, Laterallobus mit apical gerich- 
tetem Secundarlobus; oder Laterallobus sehr breit und sein Vorderrand 
etwas apical vorgezogen," and takes Pseudothelphusa macropa Milne- 
Edwards, 1853, as the "Typus subgeneris." He also assigns his Hypo- 
lobocerus guenteri to Neostrengeria. The spiny armature of the gono- 
podal funnel margin, however, enables one to separate my Phyllothel- 
phusa from Neostrengeria at first glance. In H. guenteri the carpal 
spine of the cheHpeds is prominent and acute, and is situated at about 
the middle, or just behind the middle, of the carpal margin on which the 
spine sits; in H. niceforoi, on the other hand, there is no sharp carpal 
spine, but just a low, blunt, tuberculiform projection situated within 
a third of the length of the carpus from its articulation with the palm. 

For convenience in comparing the gonopods of the several species dis- 
cussed, figures of the gonopods and carapace outhne of a specimen of 
H. (P.) Undigiana about the size of the Paris holotype are included in 
this account (Fig. 5, f-h). The specimen from which the drawings were 



108 Proceedings of the Biological Society of Washington 

made and which lacks locality data was received in exchange from the 
Museum d'Histoire Naturelle, Paris. 

Conclusions 

Variation in American freshwater crabs seems not to have 
received the attention it deserves in authors' remarks upon their 
species, at least among the Trichodactylidae. In some species 
there are noticeable differences in external form (gonopods 
aside) between male and female, and between specimens of 
the same sex, for that matter, from young to old, and from 
small to large individuals — all vary in some degree in outline, 
spination, "sculpture," and convexity of the carapace. The gono- 
pods alone seem to remain the one stable character upon which 
the specific distinctions in these brachyurans can be based. 

Much the same situation obtains in the American crayfish 
genus Procambarus, which is being intensively studied by Dr. 
Horton H. Hobbs, Jr., Smithsonian Institution senior zoologist. 
He tells me that in this genus the mature male gonopods afford 
the only reliable character by means of which its species can 
be distinguished from one another. Indeed, some are so much 
ahke that it is impossible to tell them apart without recourse 
to the gonopods. These, moreover, seem not to vary to any 
appreciable extent from the norm, no matter what the external 
morphology of the species may otherwise exhibit in the way of 
similarity or variability. 

These observations are certainly applicable to some of the 
freshwater Brachyura of the Americas. For their precise identi- 
fication, as with the North American procambarids, figures 
of the gonopods are a most essential part of every specific 
diagnosis or description. The brief descriptions and diagnoses 
of some preliminary announcements of new subspecies, spe- 
cies, subgenera, and even genera can be of but limited value 
in characterizing such taxa in the absence of illustration of the 
gonopods if recourse cannot be had to the types, if mature 
males, or to specimens that have become established as vaHd 
male representatives of the female types. 

In this connection, I wish to commend several well illus- 
trated reports concerned with the Latin American freshwater 



Colombian freshwater crab notes 109 

crabs, for in them are figured the gonopods of most of the 
species with which the authors dealt at the time: Smalley, 
1964a, b, c; Rodriguez, 1966a, b; Bott, 1956. To complete the 
characterization of Dr. Smalley's subgenus Achlidon [1964b], 
of which Pseudothelphusa agrestis Rathbun, 1898, is the type, 
the opportunity is here taken to publish a more detailed draw- 
ing, prepared some time ago, of the gonopods of that species 
than accompanied its original description (Fig. 6, a-d). 

Long before Dr. Pretzmann described V. (V.) piriformis my 
interest in variation, as observed in these Colombian fresh- 
water crabs, had been aroused by the differences noted in the 
general appearance of their carapaces, and, more particularly, 
in the armature of their anterolateral margins. This armature 
may vary from the very clear-cut, sharp spines of the smallest 
male of V, (V.) piriformis (?), which lacks abdomen and gono- 
pods (Fig. 3d), to one in which the spining has become almost 
wholly obsolescent, as in the largest male (No. 23) (Fig. 3e). 

In the largest male the extra-orbital angle is no more than 
tuberculiform, if it can be so described; following this, and 
corresponding each to the relative position of the spines pres- 
ent in the smallest male, the lateral margin of this specimen 
has mere blunt-angled notches, diminishing from front to back 
to the last, or fifth, one, which is so slight that it does not en- 
gage one's fingernail drawn along the smooth and no more 
than undulating margin at this level of the carapace. 

This progressive reduction in size and prominence or ob- 
solescence of the lateral spines seems to go hand in hand with 
the increase in size (and age?) of the males of this species and 
at least one other in the collections of the U. S. National Mu- 
seum. 

As early as 1926 Dr. Mary J. Rathbun noted on a label 
placed in a bottle of Trichodactylus (Dilocarcinus) pictus 
( <Holthuisia pictus ) from Acequia, Tucuman, Argentina ( US- 
NM 125134): "Spines of largest specimen singularly aborted." 
There were three males in the lot. The carapaces of the largest 
(44.2 X 52.6 mm) and smallest (33.5 X 39.6 mm) of these 
have been drawn in outHne (Figs. 7e and f), as well as the 
carapace of an intermediate-sized specimen (USNM 119881, 



110 Proceedings of the Biological Society of Washington 

43.6 X 50.6 mm) from Ovata, Tucuman (Fig. 7d) in order to 
show the progressive reduction in prominence of lateral spines 
that seems to accompany growth in size in this species. The 
largest of Miss Rathbuns pictus male specimens does have 
sharp-spined extra-orbital angles, and the fifth or most posterior 
"angle" on the right lateral margin of the carapace, counting 
from the extra-orbital, is armed with a sharp, quite slender 
spine, adventitious perhaps at this age or size (growth stage) 
of this male. The gonopods of the Ovata individual are figured 
(Fig. 7, a-c). These appendages, except for relative size, are 
alike in all three. 

One wonders in how many other trichodactylid species, 
given a sufficient series, such a diminution, regression, or 
abortion of the marginal armature of the carapace occurs. With 
the females, at least in V. (V.) piriformis, the situation is quite 
otherwise. There is no reduction in the relative size of the 
lateral spines from the small to the large specimens; indeed, 
the very largest female in the collection (No. 22, 54 X 59.2 
mm) is very conspicuously sharp-spined! 

Acknowledgments 

First of all, I thank Brother Niceforo Maria for these very 
interesting crustaceans now for the most part accounted for. 
I also greatly appreciate his patiently waiting so long for the 
identifications, some of which were worked over by Dr. Ger- 
hard Pretzmann and published by him elsewhere (1965- 
1968). 

I am indebted to my colleagues. Dr. Fenner A. Chace, Jr. 
and Dr. Horton H. Hobbs, Jr., for friendly and truly helpful 
advice, which, if it had been followed more closely, would 
have resulted in a better paper; to Mrs. Aime Awl, formerly 
staff artist to the Department of Zoology for the illustrations; 
to Miss Lucile McCain for the excellence of her indispensable 
editorial assistance; and to the U. S. National Museum for fa- 
cilities so generously and freely furnished me over the years. 
May Brother Niceforo be encouraged by our efforts to continue 
his interest in making the carcinological fauna of his homeland 
more completely known. 



Colombian freshwater crab notes 111 

Literature Cited* 

BoTT, Richard. 1956. Decapoden (Crustacea) aus El Salvador. 3. 
Siisswasserkrabben (Pseudothelphusa). Senck. biol. 37 (3/4): 
229-242, pis. 28-35. 

Pretzmann, Gerhard. 1965. "Vorlaufiger Bericht iiber die Familie 
Pseudothelphusidae." Sonderabdruck aus dem Anzeiger der 
math.-naturw. Klasse der Osterreichischen Akaderoie der 
Wissenschaften, Jahrgang 1965, Nr. 1, pp. 1-10. (Sitzung 
der mathematisch-naturwissenschaftlichen Klasse vom 14. 
Janner 1965.) 

. 1967. liber einige siidamerikanische Siisswasserkrabben 

(Pseudothelphusidae). Vorlaufige Mitteilung. Entomologis- 
ches Nachrichtenblatt, 14. Jahrgang, Nr. 2, Feber 1967, pp. 
23-26. 

. 1968a. Neue siidamerikanische Siisswasserkrabben (Vor- 
laufige Mitteilung). Entomologisches Nachrichtenblatt 15. 
Jahrgang, Sonderheft, Feber 1968, pp. 1-15. 

. 1968b. Weitere neue siidamerikanische Siisswasserkrabben 

(Vorlaufige Mitteilung). Entomologisches Nachrichtenblatt, 
15. Jahrgang, 2. Sonderheft, 19. April 1968, pp. 1-8. 

. 1968c. Die Familie Trichodactylidae (Milne-Edwards, 

1853) Smith 1870 (Vorlaufige Mitteilung). Entomologisches 
Nachrichtenblatt, 15. Jahrgang, Nr. 7-8, 14. Juni 1968, pp. 
70-76. 

Rodriguez, Gilberto. 1966a. The freshwater crabs of the genus Pseu- 
dothelphusa from northern Venezuela and Trinidad. Zool. 
Meded. Leiden, 41(6): 111-135, text figs. 1-10, pis. 1-7. 

. 1966b. Three new species of Pseudothelphusa from Vene- 
zuela. Zool. Meded. Leiden, 41(19): 259-267, text figs. 1-4, 
pis. 1-3. 

Smalley, Alfred E. 1964a. A terminology for the gonopods of the 
American river crabs. Syst. Zool., 13(1): 28-31, text fig. 1, 
A-H. 

. 1964b. The river crabs of Costa Rica, and the subfamilies 

of the Pseudothelphusidae. Tulane Studies in Zoology, 12 ( 1 ) : 
5-13, text figs. 1-17. 

. 1964c. Two new fresh-water crabs from Nicaragua. Ann. 

Mag. Nat. Hist. (13)7: 651-656, text figs. 1-8. 



* All other pertinent references will be fotind in the papers here listed. 



112 Proceedings of the Biological Society of Washington 



'3 



QH 

1 

B4X 


.1. 82, 


pp. 113-128 




29 May, 


1969 


HV\ 






PROCEEDINGS 
OF THE 










BIOLOGICAL SOCIETY OF WASHINGTON 





INDIAN OCEAN KINORHYNCHA 

2. NEOCENTROPHYIDAE, A NEW HOMALORHAGID 

FAMILY 

By Robert P. Higgins 
Smithsonian Institution, Washington, D. C. 

Perhaps the most criticized of Zelinka's kinorhynch species 
are Pycnophyes quadridentatus Zelinka, 1928, and P. flagel- 
latus Zelinka, 1928, each described from a single specimen. 
Both specimens were dredged from the Gulf of Baja ( Naples ) 
along with "several" P. communis Zelinka, 1928, Echinoderes 
dujardini Claparede, 1863, and several juvenile stages. 

The aberrant morphology of P. quadridentatus and P. flagel- 
latus combined with the problem of a single specimen of each 
species must have caused Zelinka considerable concern. Su- 
perficially the specimens were homalorhagid, but close inspec- 
tion revealed an undeniable resemblance to the cyclorhagid 
taxa. 

Possibly because of justification of these new species ZeHnka 
(1928, p. 136) reversed his prior opinion that "paedogenesis" 
occurred in the homalorhagids. Nonetheless, paedogenic char- 
acters were manifest in the morphology of these strange new 
species which remained unchallenged for nearly 20 years. 

In 1947, Nyholm (p. 36) reported: "that paedogenetic Hy- 
alophyes forms, when moulting, discharge eggs in the old 
integuments" and (p. 35) "that Hyalophyes forms — can be- 
come sexually mature but they continue to grow and not reach 
the imago until they change to Pycnophyes forms." I concur 
with Nyholm in that it does appear that the last juvenile stage 
of Pycnophyes may become sexually mature, but in such cases 
there is not a marked degree of morphological difference be- 
tween this form and the normally developed adult. In cases 

7— Prog. Biol. Soc. Wash., Vol. 82, 1969 (113) 



Nw2bf969J 



114 Proceedings of the Biological Society of Washington 

where the last molt is omitted the total length seems to be the 
most striking difference between the matured juvenile and the 
adult which has undergone the final molt, and the length char- 
acteristics of the two forms overlap in their normal curves. 

Soon after Nyholm began and finished his work on kino- 
rhynchs, Lang (1949), in addition to describing new kino- 
rhynchs from the Swedish Antarctic Expedition, 1901-1903, 
critically examined the status of kinorhynch phylogeny and 
taxonomic characters and, upon reviewing the homalorhagids 
(p. 12) stated: "The isolated position of quadridentatus, flagel- 
latus and echinoderoides in the groups of Zelinka as well as of 
Remane makes one extremely skeptical about them. And a 
study of Zelinka's reports on them — the three species have all 
been established by him — leads to the conviction that they 
lack any right of existence." 

The report which follows is the third (Higgins, 1968a, b) in 
a series on the extensive collections of kinorhynchs I made 
along the coasts of India, Kenya, and the island of Nosy Be, 
Malagasy Republic in 1964. The methods are identical with 
those outlined in the second paper of the series (Higgins, 
1968b) which discusses the cyclorhagid families Semnoderidae 
and Centroderidae. 

In this paper I shall resurrect the Lang-Zelinka dispute 
which has been mentioned above because I share Zelinka's 
experience of finding too few specimens — specimens which, 
as did the Zelinka collection from the Gulf of Baja, clearly re- 
veal morphology intermediate, in several characteristics, be- 
tween the cyclorhagid and homalorhagid taxa. 

Acknowledgments: I would like to thank Dr. T. S. Satyana- 
rayna Rao, Liaison Officer for the U. S. Program in Biology, 
International Indian Ocean Expedition, for his extensive help 
in arranging my research in India; Dr. A. H. Humes, Boston 
University, for his assistance in the capacity of Chief Scientist, 
IIOE, at Nosy Be, Malagasy Republic; Dr. P. N. Ganapati 
and the staff of the Department of Biology, Andhra Univer- 
sity, Waltair, India and M. Angot, Centre d'Oceanographie et 
des Peches, Nosy Be, Malagasy Republic, for making the facil- 
ities of their respective institutions available to me and pro- 



Indian Ocean Kinorhyncha 115 

viding assistance in the collecting activities; and to Drs. H. L. 
Sanders and R. R. Hessler, Woods Hole Oceanographic Insti- 
tution, my companions in this portion of the IIOE. 

This study was made possible by the National Science 
Foundation's support of the U. S. Program in Biology, IIOE, 
and grant GB 3069 which is gratefully acknowledged. This 
manuscript was written while the author was Acting Resident 
Systematist with the Systematics-Ecology Program, Marine 
Biological Laboratory, Woods Hole, Massachusetts. 

The following key is amended from the prior pages in this 
series (Higgins, 1968b) to include the new genus and family 
whose descriptions will follow. 

Key to the Adults of the Kinorhynch Genera 

1. First trunk segment entire, bivalvate with or without midventral 
and middorsal plates; round or oval in cross section; lateral, 
dorsal and terminal spines usually well developed; with 14-16 

placids (neck plates) when present order Cyclorhagida 

First trunk segment with single, strongly arched dorsal plate, 
single midventral plate and two lateroventral plates, or with 
ventral plates partially or entirely fused to form single, broad 
ventral plate; articulated (movable) trunk spines, if present, 
restricted to lateroterminal spines, occasionally with distinct 
spinose extensions of middorsal and lateral tergal borders, with 
6-8 placids order Homalorhagida 2 

2(1). With midterminal spine and distinct spinose extensions of mid- 
dorsal and lateral tergal borders; with 7 placids 

Neocentrophyidae, n. f., Neocentrophyes, n. g. 

Without midterminal spine, with 6 or 8 placids 3 

3(2). With lateroterminal spines Pycnophyes 

Without lateroterminal spines Trachydemus 

4(1). First trunk segment entire, without midventral plate 

suborder Cyclorhagae 5 



First trunk segment bivalvate, and/or with midventral plate; if 
not bivalvate, midventral plates on segments 3-8 8 

5(4). First and second trunk segments entire; midterminal spine ab- 
sent family Echinoderidae, Echinoderes 

First trunk segment entire, second trunk segment with two 

lateroventral plates; midterminal spine present 

family Centroderidae 6 

6(5). Elongate spines extending from posterolateral margins of first 
trunk segment; lateroterminal accessory spines present in addi- 
tion to lateroterminal spines 7 



SMimsoNiAN MAY 2 9 1969 

iNsinuTior 



116 Proceedings of the Biological Society of Washington 

Elongate spines not present on posterolateral margins of first 
segment; lateroterminal accesory spines absent Condyloderes 

7(6). Lateral spines on most trunk segments; few lateral accessory 

spines present Campyloderes 

Lateral spines on few trunk segments, without lateral accessory 
spines Centroderes 

8(4). Midventral plate on segments 3-8, tapering in posterior pro- 
gression; lateroventral articulation zone weakly developed; mid- 
dorsal spine absent on segments 3, 7, 9, and 12 

. suborder Cryptorhagae, family Cateriidae, Catena 



Midventral plate, if present, restricted to first trunk segment, first 
trunk segment with bilateral plates; lateroventral articulation 

zone obvious; middorsal spines on all tnmk segments 

suborder Conchorhagae, family Semnoderidae 9 

9(8). Midventral and middorsal plate present on first trunk segment; 

with well-developed placids Sphenoderes 

Midventral and middorsal plate absent; without distinct placids 
Semnoderes 

Order Homalorhagida (Zelinka, 1896) Chitwood, 1958 
Homalorhagae Zelinka, 1896, p. 198. 
Homalorhaga Gerlach, 1956, p. 129. 
Homalorhagea Chitwood, 1958, p. 942. 

Definition: Kinorhyncha with second segment (neck) consisting of 
6, 7, or 8 well-formed placids (neck plates); trunk segments flattened 
ventrally, vaulted dorsally, triangular in cross section; protonephridia 
without ligaments; gonopores ventrolateral on segment 13 or lateral 
between segments 12-13; pharynx musculature tripartite; oblique mus- 
cles absent; armor joints, if present, consisting of dorsal acetabulum and 
ventral condyle; with intersegmental processes along middorsal ridge or 
non articulate spinous processes both middorsally and laterally on tergal 
plates, if the latter, then with well-developed perispinal setae on either 
side of spinous processes; occasionally with midterminal spine in early 
postembryonic stages or adult; surface of trunk segments with minute 
pores or denticulate, hair like processes. 

Neocentrophyidae new family 
Definition: Homalorhagida with second segment (neck) consisting of 
seven well-formed placids (four dorsal and three ventral); mouth cone 
with four short, thin, unjointed styles and five longer, two-jointed styles, 
all with pectinate basal area; six(?) rows of scalids present, first row 
with single elongate seta centered on basal plate, bordered by pectinate 
fringe; 14 trichoscalids present in posterior scalid row; first trunk seg- 
ment not divided into three ventral plates, or with plate formation evi- 
denced only at anterior margin; middorsal and lateral, non articulate 
spinous processes on all tergal plates; Mdth well-developed perispinal 
setae on either side of most spinous processes; midterminal spine pres- 



Indian Ocean Kinorhyncha 117 

ent, short, located between median terminal extensions of segment 13 so 
as to appear at the base of a notch in the terminal segment; armor 
joints absent or weakly developed; surface of trunk segments with dentic- 
ulate hairlike processes; gonopores lateral between segments 12-13. 

Neocentrophyes new genus 

Type species: Neocentrophyes intermedius new species. 
Definition: As in family. 

Neocentrophyes intermedius new species 
Figures 1-3 

Diagnosis: Neocentrophyes with nonsetate trichoscalids; with ventral 
perispinal seta of lateral spinous process missing on segments 4, 6, 8, 
and 12; with middorsal flagelloid spinous process extending from pos- 
terior margin of segments 12 and 13, other middorsal spinous processes 
reaching maximum length of 30^ at segment 7. 

Holotype: Adult male, TL 650^; Ambatozavavy Bay, Nosy Be, Mala- 
gasy Republic (13° 21.5' S, 48° 19.5' E); 1-2 m; gray-brown sandy mud; 
15 April 1964; col. R. P. Higgins (RH44.81); USNM 37993. 

Paratypes: 4 adult males, TL 490-608/>i, 1 juvenile, TL 352/*; as holo- 
type (RH44 series); 1 adult male, TL 554/x; Anse des Antalaotra, Nosy 
Be, Malagasy Republic; 1-2 m; muddy sand; 16 April 1964; col. R. P. 
Higgins ( RH45 series ) ; 2 adult males, TL 570-577/*; Ampasindava Bay, 
Nosy Be, Malagasy RepubHc; 18 m; gray mud; 23 April 1964; col. R. P. 
Higgins (RH51 series); RH44, RH45, and RH51 series retained in 
author's collection. 

Description: Holotypic male, TL 650/*; MSW-5 170/*; SW 120/*; 
SW/TL 0.19. 

First segment (head) retracted. The following description of the head 
is based on a paratypic male, TL 608/t, RH44.78 (Fig. 3): Mouth cone 
with five two-jointed styles, 80/t, and four non jointed styles, 45/i, all 
with pectinate basal plate; six(?) rows of scalids, 10 spinoscalids of 
first row blunt-tipped, ca. 105/jl, with pectinate basal plate and single 
elongate seta, ca. 40/*; second through fifth rows with pointed spino- 
scalids, ca. 75/t, 73/*, 57/t, and 52/*; posteriormost row with 14 non- 
setate trichoscalids, 45/t. 

Second segment (neck) with four dorsal placids (Fig. 1), medial 
placids 40/* wide, lateral placids 25/* wide; with three ventral placids 
(Fig. 2), midventral placid 64/i wide, lateral placids ca. 30/* wide. 

Third segment (first trunk segment) with anterolateral margins pro- 
jecting slightly as horn-Uke processes; anterior margin of dorsal and 
ventral plates even; muscle scars barely evidenced posterolaterally on 
dorsal plate and anterolateral on ventral plate; surface of plates with 
scattered openings, probably sensory pores; lateral tergal margin extended 
posteriorly as spinous process, ca. 27/*, with prominent perispinal setae, 
ca. 20/t, adjacent to base, one on ventral plate, one dorsal to lateral 
spinous process; middorsal ridge extending posteriorly to form short 



118 Proceedings of the Biological Society of Washington 





Figures 1-2. Neocentrophyes intermedius n. sp., adult male, holo- 
tj^e, TL 650/tt, neck and trunk segments: 1, dorsal view; 2, ventral view. 



spinous process, ca. ISfi, also with perispinal setae, ca. 20(0,, adjacent 
to base; posterior margins of dorsal and ventral plates thin, finely striate. 
Fourth segment with lateroventral plates, indistinctly articulating 
laterally with dorsal plate, typical homalorhagid armor joints absent or 
poorly developed; dorsal perispinal seta present near base of lateral 



Indian Ocean Kinorhyncha 



119 



T 

50|j 

i 




Figure 3. Neocentrophyes intermedius n. sp., adult male, paratype, 
TL 608fi, diagrammatic perspective, lateral half of head segment showing 
portion of mouth cone and spinoscalid-trichoscaHd series. 



120 Proceedings of the Biological Society of Washington 

spinous process, ventral perispinal seta absent; four sensory pits near 
anterior margin and three near posterior margin of each ventral plate and 
left and right halves of dorsal plate; otherwise similar to third segment. 

Segments 5-11 similar to fourth, middorsal spinous processes reaching 
maximum length at segment 7, D-5 19/*, D-6 28/x, D-7 30/i, D-8 24fi, 
D-9 27 fi, D-10 20/A, D-11 28/*; lateral spinous processes slightly longer, 
L-5 31/*, L-6 32/*, L-7 35/*, L-8 33/*, L-9 28/*, L-10 26/*, L-11 26/*; 
ventral perispinal seta of lateral spinous process absent on segments 6 
and 8. 

Segment 12 tapering posteriorly; with fewer sensory pits dorsally; 
lateral spinous process not extending beyond striate border of segment, 
ventral perispinal seta of lateral spinous process absent; middorsal 
spinous process thin, flagelloid, 50/*. 

Segment 13 bifurcate, similar to Echinoderes; deeply incised along 
midline to accommodate short, midterminal spine, 29/t, nearly extending 
to margin of segment; with thin, flagelloid middorsal spinous process, 
60/*; penis spine present anterolaterally, 48/t, second penis spine slightly 
posterior on lateral margin, 38/*; terminal margin of dorsal plate not 
extending to terminal margin of ventral plate; bulbose protrusion at 
lateroterminal junction of dorsal and ventral plates, 8/*. 

Measurements (/*) and indices for selected specimens of Neocentro- 
phyes intermedius n. sp. are as follows: 





Juvenile 
RH44.80 


Adult d Adult d 
RH44.82 RH44.76 


Adult d 
RH51.9 


Adult d 

USNM 
37994 


Adult d 

USNM 

Holotype 


Adult 
Range 


TL 


352 


490 


527 


577 


608 


650' 


490- 650 


SW 


96 


114 


112 


120 


120 


120 


112- 120 


SW/TL 


0.27 


0.23 


0.23 


0.21 


0.20 


0.19 


0.19-0.23 


MSW-5 


130 


154 


168 


160 


168 


170 


154- 170 


MTS 


14 


16 


14 


14 


19 


20 


14- 20 


MTS/TL 


0.040 


0.033 


0.027 


0.024 


0.031 


0.031 


0.024-0.033 



Discussion: Neocentrophyes intermedius n. sp. resembles Pycnophyes 
flagellatus in size (TL 595/* for the latter species), in its spinous proc- 
esses and perispinal setae, in the denticulate hair-hke processes, in the 
presence of a midventral placid, and most acutely, in the presence of the 
middorsal flagelloid spinous process on the twelfth segment and similar 
process on the thirteenth segment. The new species differs from P. 
flagellatus by the former's lack of lateroterminal spines, by its lack of a 
broadly rounded thirteenth tergal plate, and apparently thickened but 
not articulate midventral and lateral plates on the ventral surface of the 
first trunk segment. Additionally, P. flagellatus apparently lacks(?) a 



Indian Ocean Kinarhyncha 121 

midterminal spine and has some delineation of the typical homalorhagid 
plate formation on the ventral surface of segment 3. 

The normal attention to detail in the majority of Zelinka's species 
descriptions and illustrations deteriorates with P. flagellatus. The char- 
acter of the mouth cone, by itself, in Neocentrophyes intermedius, n. 
sp. is of sufficient magnitude to separate it generically from Pycnophyes. 
Nothing is known of the mouth cone structure in P. flagellatus. 

In the case of P. quadridentatus, N. intermedius, n. sp. differs by 
its lack of lateroterminal spines, by its flagelloid thirteenth middorsal 
spinous process, and by its lack of marginal plate formation on the first 
trunk segment. In both P. quadridentatus and P. flagellatus the mid- 
ventral placid is no wider than the lateral placids, but in N. intermedius, 
n. sp. this placid is twice the width of those adjacent to it. 

Considering the morphology of the juvenile specimen of N. intermedius, 
n. sp., TL S52fi, which lacks the flagellar spinous processes of the 
adult, it seems possible that P. quadridentatus may be a juvenile stage 
of P. flagellatus. Only by intensive study of the kinorhynch populations 
where these presently discussed species occur can the matter be re- 
solved. 

Neocentrophyes satyai new species 
Figures 4-5 

Diagnosis: Neocentrophyes with setate trichoscalids, with both dorsal 
and ventral perispinal setae missing at base of lateral spinous process 
of segment 4; without middorsal flagelloid spinous process on terminal 
segments, other spinous processes reaching maximum length of 52^1 at 
segment 6. 

Holotype: Adult female, TL 537^; ca. 10 km offshore E of Visak- 
hapatnam (Bay of Bengal), India; 40 m; brown sandy mud; 26 March 
1964; col. R. P. Higgins (RH37.87); USNM 37995. 

Paratypes: Adult female, TL 445/* and juvenile, TL 430/a; as holotype; 
(RH37 series, retained in author's collection). 

Description: Holotypic female, TL 537/*; MSW-8 146/t; SW 124/i; 
SW/TL 0.23. 

First segment (head) retracted. Second segment not visible except 
for retracted outline which shows seven placids as in generic definition. 

Third segment (first trunk segment) with anterolateral margins pro- 
jecting slightly as horn-like processes; anterior margin of dorsal and 
ventral plates even; large muscle scars discernible only on ventral plate, 
centered midway between anterior and posterior margins; sensory pores 
not well-evidenced; lateroventral margin of dorsal plate extended into 
spinous process, ca. 37/*, with prominent perispinal setae, ca. 20/*, 
adjacent to base, one on ventral plate, one dorsal to lateral spinous 
process; middorsal ridge extending posteriorly to form spinous process, 
35/*, with perispinal setae, ca. IS/t, adjacent to base; with posterior 
margins of dorsal and ventral plates thin, finely striate. 

Fourth segment with lateroventral plates indistinctly articulating 



122 Proceedings of the Biological Society of Washington 

I 

i 



100y 





Figures 4-5. Neocentrophyes satyai n, sp., adult female, holotype 
TL 537/A, trunk segments: 4, dorsal view; 5, ventral view. 



laterally with dorsal plate, typical homalorhagid armor joints absent or 
poorly developed; without lateral perispinal setae; no sensory pits noted 
(possibly obscured by condition of specimen). 

Segments 5-11 similar to fourth but with lateral perispinal setae; 
middorsal spinous processes reaching maximum length at segment 6, 
D-5 40At, D-6 52m, D-7 52/*, D-8 52/*, D-9 33/*, D-10 28/*, D-11 30/*; 



Indian Ocean Kinorhyncha 123 

lateral spinous processes slightly shorter, L-5 42//., L-6 37/*, L-7 40/x, 
L-8 41/., L-9 41/i, L-10 31/t, L-11 25/.. 

Segment 13 bifurcated as in Neocentrophyes intermedius n. sp.; deeply 
incised along midline to accommodate short midterminal spine, ca. 20/*, 
middorsal spinous process barely suggested; margin of dorsal plate not 
extending to terminal margin of ventral plates; bulbose protrusion at 
lateroterminal junction of dorsal and ventral plates, 15/.. 

Measurements (/.) and indices for specimens of Neocentrophyes satyai 
n. sp. are as follows: 





Juvenile 
RH37.27 


Adult 9 
RH37.88 


Adult 9 

USNM 37995 

Holotype 


TL 


430 


445 


537 


SW 


102 


104 


124 


SW/TL 


0.24 


0.23 


0.23 


MSW-8 


124 


140 


146 


MTS 


18 


18 


20 


MTS/TL 


0.043 


0.040 


0.037 



Etymology: This species is named in honor of the Liaison Officer for 
the U. S. Program in Biology, IIOE, Dr. T. S. Satyanarayna Rao— 
respectfully and affectionately called "Satya" by his many friends. 

Discussion: Neocentrophyes satyai n. sp. differs from N. intermedius 
by the former's lack of the flagelloid middorsal spinous process on seg- 
ments 12 and 13, lack of lateral perispinal setae on segment 4 and pres- 
ence of both dorsal and ventral perispinal setae on the remainder of the 
lateral spinous processes of the trunk segments, by its significantly 
shorter (13/.) middorsal spinous process on segment 3 and much longer 
processes on segments 4-8 in particular. Characteristics such as the more 
posterior position of the lateral perispinal setae of segment 3 (Figs. 4-5), 
the presence of a small seta near each lateroterminal protuberance, the 
outline of the terminal border of both dorsal and ventral plates, and the 
MTS/TL index may be of taxonomic importance. 

The presence of a lateroterminal spine in both Pycnophyes flagellatus 
and P. quadridentatus readily separates these two species from N. satyai 
n. sp. 

Neocentrophyes satyai n. sp. is known only from two adult females 
just as N. intermedius n. sp. is known only from several adult males. 
Although nothing can be stated about the sexual dimorphism of either of 
the new species, based upon my experience, I would not expect to find 
sexual dimorphism accounting for the differences between N. intermedius 
n. sp. of the West Coast of Madagascar and N. satyai n. sp. from the East 
Coast of India. 



124 Proceedings of the Biological Society of Washington 

The Evolutionary Significance of the Neocentrophyidae 

Zelinka's aberrant species, Pycnophyes flagellatus, P. quadri- 
dentatus and P. echinoderoides, each based on a single speci- 
men, have been banished from existence by Lang, 1949 as 
noted in the introduction of this paper. Although I, too, would 
exclude P. echinoderoides at this time, and have serious doubts 
as to the taxonomic validity of both P. flagellatus and P. quad- 
ridentatus (the latter possibly a juvenile stage of the former), 
I shall renew the Lang-Zelinka dispute. 

Indeed, I shall take up the argument for the existence of 
Zelinka's disputed taxa not merely because of the present dis- 
covery of two additional species which, by Lang's reasoning, 
would be explained as deformations or retarded development, 
but because of the rapidly accumulating evidence that heter- 
ochrony ( =paedogenesis ) has been an important factor in 
the establishment of several uncommon kinorhynch taxa. 

The notation by Nyholm ( 1947 ) in his studies of the Kino- 
rhyncha, that "paedogenesis" may occur in Pycnophyes is cer- 
tainly substantiated by my own studies (Higgins, 1962). Yet, 
Lang (1949) is correct in raising many questions basic to this 
feature of kinorhynch biology. Perhaps the only contribution 
that can be added to Nyholm's work is the observation that 
when the fifth of a normally six-instar series in Pycnophyes as- 
sumes adult characters without the final molt, this individual 
is not unrecognizable as conspecific with the more normally 
developed adult of the species in question. The recognition, 
however, of earlier stages is an entirely different matter. 

Probably the most aberrant and obviously neotenic kino- 
rhynch taxon to be described so far is Catena ( Gerlach, 1956 ) . 
In my recent discussion (Higgins, 1968a) of this mesopsam- 
mic taxon I mentioned the following series of juvenile char- 
acters among the Kinorhyncha: 1) a poorly developed cuticle, 
usually hyaline rather than chitin "tanned"; 2) spinous to 
denticulate cuticular marking rather than hairs or pores which 
communicate with the underlying cells; 3) incomplete or 
poorly developed segmental or plate articulation, particularly 
lateroventrally; 4) presence of midterminal spine, laterotermi- 
nal and lateroterminal accessory spines, and extensive mid- 



Indian Ocean Kinorhyncha 125 

dorsal and lateral spines; 5) absence of sexually dimorphic 
characteristics, including various spines, certain adhesive 
tubes, and apparent absence of functional gonopores. 

Recently described new genera such as Condyloderes Hig- 
gins, 1968b and Sphenoderes Higgins, 1968b reflect the grow- 
ing evidence that the ontogeny of the better known taxa such 
as Echinoderes and Pycnophyes is reflected in the adults of 
less common taxa. Certainly the presence of the genus Neo- 
centrophyes is well within the expected evolutionary limits of 
the phylum. 

Assuming the validity of the two most common genera of 
Homalorhagida, ie. Pycnophyes and Trachydemus, the former 
differing from the latter by its possession of lateroterminal 
spines and recognizably distinct postembryonic stages, and 
assuming the heterochronic potential of the various taxa now 
known, it then follows certain juvenile characters might estab- 
lish themselves in a taxon closely related to the more "adult" 
genus Pycnophyes and, similarly, the genus Trachydemus. 
Indeed, this is strongly evidenced by Neocentrophyes which, 
by its lack of lateroterminal spines, suggests a strong relation- 
ship to Trachydemus. A similar relationship exists between 
Pycnophyes and Zelinka's P. flagellatus and P. quadridentatus 
as suggested in their being assigned to this genus by Zelinka 
on the basis of their lateroterminal spines. 

Following his declaration that P. flagellatus, P. quadri- 
dentatus and P. echinoderoides "lack any right of existence," 
Lang (1949, p. 12) continues: "Each of them has been ob- 
tained once only. They all have in common the weak cuticu- 
larization which may indicate their having moulted but re- 
cently. Echinoderoides was found also still lying in the Hya- 
lophyes skin. And Zelinka ( I.e., p. 218 et al. ) has pointed out 
himself that the Echinoderida directly after their metamor- 
phosis' look quite different from older imagines." 

The unfortunate possibility that representatives of certain 
taxa may be collected "once only" either because of their 
normally low proportion within the community sampled or 
because they are not normally a component of the sampled 
area, or for any reason, does not justify their "non-existence." 



126 Proceedings of the Biological Society of Washington 

Weak cuticularization is now a well-established character re- 
tained by such neotenic taxa as the Cryptorhagae. The present 
evidence does not allow for comment on the validity of all 
three of the species criticized by Lang to any further degree 
than has already been discussed. 

Lang continues: "The extraordinary structure of the 3rd 
zonite in quadridentatus, which, incidentally was already dead 
when secured, cannot be considered a normal phenomenon. 
The impression it makes of being a defective specimen, the de- 
velopment of which has been retarded, is increased not only 
by the above mentioned shape of the armour, but also by its 
having kept such 'larval' characters as the strong lateral and 
dorsal spines and especially the long dorsal spines situated 
almost terminally on the last zonite (cf. Nyholm 1947b, p. 9)." 

The structure of the third segment (first trunk segment) of 
the taxa in question is neither unexplainable nor unexpected 
considering the morphology of this same segment in both 
Catena and Sphenoderes, neither of which was known to Lang. 
In Catena, an otherwise normal cyclorhagid morphology is 
contrasted with the distinct division of the first trunk seg- 
ment's ventral surface into a midventral plate which, depend- 
ing upon the species, may not be separated from the remain- 
ing portion of the segment for its entire length. In Catena, 
this ventral plate formation proceeds to the eighth segment 
and is accompanied by weak development of the lateroventral 
articulation zone. In Sphenoderes, the otherwise Semnoderes- 
character of the genus is marked by the presence of both a 
middorsal and midventral plate between the bilateral clam- 
shell-like closing apparatus formed by the first trunk segment. 

In each case. Catena and Sphenoderes, two otherwise cyclor- 
hagid forms ( 14 placids or their equivalent on segment 2, etc. ) 
have exhibited homalorhagid morphology by the development 
of a midventral plate on segment 3. The "larval" characters 
such as the strong lateral and dorsal spines, etc. that Lang 
refers to, and which exist in Neocentrophyes as well as the 
Zelinka species to which Lang alludes, are extremely sugges- 
tive of the postembryonic stages of the homalorhagids just as 
is the poorly developed plate formation on the third segment. 



Indian Ocean Kinorhyncha 127 

On the other hand, several previously described genera such 
as Semnoderes, Centroderes, Campyloderes, and the more 
recently described Sphenoderes and Condyloderes retain the 
juvenile character for both middorsal and lateral spines to be 
extensively developed in the adult. Neocentrophyes follows 
this same pattern as do the Zelinka species, regardless of their 
arbitrary generic assignment. 

Finally, let me comment on Lang's statement: "Flagellatus 
gives the same impression as quadridentatus. According to 
Zelinka it is a male, but that it does not represent a normally 
developed stage is indicated by its lack of penis spines, in con- 
trast to all other Homalorhagae. The lack of penis spines, like 
the lack of genital openings, is characteristic for the larvae' 
(Zelinka, I.e., p. 136, Remane, I.e., p. 331)!" 

Zelinka's illustration of P. flagellatus (Plate 14, Figures 4-5) 
clearly shows two lateral spines which he labels sbi and sb2 or 
"Seitenstacheln" which are the equivalent of the penis spines 
of Neocentrophyes intermedius (Figs. 1-2). Certainly, if one 
looks for the normal homalorhagid genital apparatus in the 
intermediate-oriented Neocentrophyes one will not find it. In- 
stead, Neocentrophyes penis spines and associated genital 
apparatus are located laterally as in the Cyclorhagida, one of 
the several characteristics which enhances the suggestion that 
this taxon is phylogenetically an intermediate through a neo- 
tenic evolutionary process not uncommon to many invertebrate 
taxa. The mature state of the gonads of the specimens de- 
scribed in this paper obviates the question of their adulthood. 

Finally, we must consider the implication that if Neocen- 
trophyes is to Trachydemus as Pycnophyes flagellatus is to 
Pycnophyes, then P. flagellatus may constitute a new generic 
category. A single specimen of homalorhagid kinorhynch taken 
from the continental slope off the coast of North Carolina by 
Dr. F. Grassle, Duke University Marine Laboratory, shows 
the essential family characteristics attributed to Neocentrophyi- 
dae. The specimen is well-sclerotized and possesses midtermi- 
nal and lateroterminal spines and unfortunately these are 
broken off near their base. The middorsal and lateral spinous 
processes, ventral aspect of the first trunk segment, perispinal 



128 Proceedings of the Biological Society of Washington 

setae, denticulate segment surface pattern, all correspond to 
the characteristics of the Neocentrophyidae. Extensive sam- 
pling may provide additional specimens of this unquestionably 
new taxon, meanwhile a ray of hope for kinorhynch beta and 
gamma taxonomy begins to appear. 

Literature Cited 

Chttwood, B. G. 1958. The classification of the Phylum Kinorhyncha. 
Proc. 15th Intern. Congr. Zool. 1958, pp. 941-943. 

Claparede, E. 1863. Beobachtungen Uber Anatomie und Entwick- 
lungsgeschichte wirbelloser Tiere an der Kiiste der Normandie 
angestellt. Wilhelm Englemann, Leipzig, pp. 1-120, pis. 1-18. 

Gerlach, S. a. 1956. Uber einen aberranten Vertreter der Kino- 
rhynchen aus dem Kiistengrundwasser. Kiel Meeresf., vol. 12, 
pp. 120-124, figs. 1-15, pis. 40-42. 

HiGGiNS, R. P. 1962. Postembryonic life history of the homalorhagid 
Kinorhyncha. Amer. Zoologist, vol. 2, p. 214 (abstract). 

. 1968a. Taxonomy and postembryonic development of the 

Cryptorhagae, a new suborder for the mesopsammic kino- 
rhynch genus Catena. Trans. Amer. Micros. Soc, vol. 87, no. 
1, pp. 21-39, figs. 1-25. 

. 1968b. Indian Ocean Kinorhyncha. 1. Condyloderes and 

Sphenoderes, new cyclorhagid genera. Smithsonian Contr. 
Zool. No. 14, in press. 

Lang, K. 1949. Echinoderida. In Odhner, J. H., ed., Further Zoo- 
logical Results of the Swedish Antarctic Expedition 1901- 
1903, vol. 4, no. 2, pp. 1-22, figs. 1-8. 

Nyholm, K-G. 1947. Studies in the Echinoderida. Ark. Zool., vol. 39 
A, No. 14, pp. 1-36, figs. 1-22, pis. 1-2. 

Zelinka, C. 1896. Demonstration von Tafeln der Echinoderes - 
Monographic Verhandl. Deutsche Zool. Ges. 6. Jahresver- 
samml. Bonn, pp. 197-190. 

. 1928. Monographie der Echinodera. Wilhelm Englemann, 

Leipzig, pp. i-iv, 1-396, pis. 1-27. 



82, pp. 129-142 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 

BIOLOGICAL INVESTIGATIONS OF THE DEEP SEA. 46. 

THE GENUS LITONOTASTER 

(ECHINODERMATA, ASTEROIDEA)' 

By Jerald a. Halpern 
Institute of Marine Sciences, University of Miami 

Two specimens of sea stars were collected by the Institute 
of Marine Sciences, University of Miami vessels R/V Gerda 
and R/V Pillsbury in the Straits of Florida and the Gulf of 
Guinea. Each represents a new species of the goniasterid 
genus Litonotaster. 

These species as well as L. intermedins, the type-species, are 
described. The diagnosis of Litonotaster is revised. 

This research was supported by National Science Founda- 
tion grant GB-4936. The operations of R/V Gerda and R/V 
Pillsbury were supported by National Science Foundation 
grant GB-7082. I thank Dr. L. P. Thomas for critically reading 
the manuscript. 

Genus Litonotaster Verrill, 1899 
Litonotaster Verrill, 1899, pp. 171-172.— Fisher, 1911, pp. 165-166. 

Diagnosis: Abactinal plates flat, very thin. All abactinal plates bearing 
more than a single marginal row, but at least some plates not completely 
covered by granules. No secondary abactinal plates. Papular areas 
narrow. Marginal plates flattened. No superambulacral ossicles; no in- 
ternal radiating ossicles. 

Discussion: This genus was erected by Verrill for the single species 
Pentagonaster intermedins Perrier. Verrill (1899: 172) separated it from 
related genera on the basis of four major characters: "few and minute 
papular pores and the very limited area on which they occur; the thin 
and small marginal plates; flexible dorsal surface of the disk; and large 
number of adambulacral spines." The use of numerous adambulacral 
furrow spines ( seven or eight ) as a generic character has since been modi- 
fied by H. L. Clark (1920: 83-85), who described a species of Litono- 
taster with four or five adambulacral furrow spines. Another important 



1 Contribution No. 1040 from the Institute of Marine Sciences, University of Miami. 

8— Proc. Biol. Soc. Wash., Vol. 82, 1969 (129) 






130 Proceedings of the Biological Society of Washington 

character of Litonotaster is the abactinal granulation. Verrill (1899: 171) 
described the abactinal plates as, "finely granulated, with two or more 
rows of granules around the edges, but with a small, central, round, 
naked area." Often, not all the plates have a central naked area, and in 
some specimens most of the plates are completely covered by granules. 

The marginal plates of Litonotaster are flattened and thin as com- 
pared to the angular, massive plates found in most goniasterid genera 
(see fig. 1). In Litonotaster intermedius and L. tumidus the superomar- 
ginals are mainly in the vertical plane, so that they appear small and in- 
conspicuous (see fig. IB). This type of marginal plate is also found in 
other goniasterid genera, e. g. Fergamaster and Tessellaster. In L. rotundi- 
granulum, n. sp. the superomarginals are mainly in the horizontal plane 
so that they appear large and conspicuous (see fig. lA). The supero- 
marginals of L. africanus, n. sp. are at an angle of about 30° from the 
horizontal. Thus, the difference between the marginal plates of the type- 
species, L. intermedius, and the marginals of the new species described 
here are moderate and not of generic importance. 

Type-species: Pentagonaster intermedius Perrier (by original designa- 
tion: Verrill, 1899, p. 171). 

Key to the species of Litonotaster 

1. Actinal plates covered by conical spinules 2 

Actinal plates covered by hemispherical 

granules rotundigranulum 

2. Actinal spinules two to three times as high as wide tumidus 

Actinal spinules about as high as wide 3 

3. Many abactinal plates bearing pedicellariae africanus 

No abactinal pedicellariae present intermedius 

Litonotaster intermedius ( Perrier, 1884 ) 
Figures IB, 2, 3 

Pentagonaster intermedius Perrier, 1884, p. 243, pi. 5, figs. 5, 6. 

non Pentagonaster intermedius: Alcock, 1893, p. 90. 

Litonotaster intermedius: Verrill, 1899, pp. 172-173, pi. 28, figs. 5, 5a, 

5b.— H. L. Clark, 1920, p. 85; 1941, pp. 43-44.— Madsen, 1951, p. 

88.— A. H. Clark, 1954, p. 375. 

Material studied: 28°00'N, 87°42'W, 2685 m, Albatross sta. 2379, 2 
March 1885, USNM 18443, 1 spec— 26°34'N, 90°31'W, 2379 m, M/V 
Oregon sta. 2571, 27 July 1959, 1 spec— 26°34'N, 89°53'W, 2654 m, 
M/V Oregon sta. 2574, 28 July 1959, 2 spec. 

Diagnosis: Abactinal plates covered by small, rounded, closely crowded 
granules; small central area of some plates bare. No abactinal pedicel- 
lariae. Actinal plates covered by short, conical spinules, about as high 
as wide. Adambulacrals bearing six to eight furrow spines. 

Description: Five arms. R = 32 mm; r = 12 mm; R/r = 2.7. General 
form stellate; disk inflated, dorsal integument thin, flexible; interbrachial 
arcs rounded. 



The genus Litonotaster 



131 




B 



Fig. 1. Lateral view of marginal plates (diagrammatic). Stippling 
indicates external surface; s, superomarginal plate; i, inferomarginal plate. 
A, Litonotaster rotundigranulum, n. sp.; B, L. intermedius; C. Rosaster 
alexandri (illustrating angular marginal plates found in most goniasterid 
genera. ) 

Abactinal plates small, flat, irregularly round, closely crowded; covered 
by small, rounded, closely crowded granules. Small central area of some 
plates bare. No abactinal pedicellariae. Interradial areas large. Papular 
areas small, inconspicuous; restricted to narrow bands running from 
bases of arms to center of disk; no papulae in center of disk. Six papular 
pores surrounding each plate, each pore containing a single papula. 

Twenty marginal plates in each series corresponding throughout. 
Superomarginals slightly longer than wide; one or two pairs in contact 
medially. Each plate surrounded by row of granules similar to those on 
abactinals. Superomarginals in interbrachial arc having lower two-thirds 
covered by coarse, rounded granules, about twice as large as peripheral 
granules; upper third naked or with several scattered granules. Naked 
area becoming larger distally, so that last three or four plates bearing 
only peripheral row of granules. In interbrachial arcs lateral angle of 
each plate depressed; plates flattened, mainly in vertical plane (see fig. 
IB). Plates becoming more angular distally. Inferomarginal plates square 
in interbrachial arc, becoming longer than wide distally; covered by short, 
conical spinules. Some plates bearing two or three valved, short, excavate 
pedicellariae; valves as high as wide. Terminal plate heart-shaped; distal 
end notched, proximal end truncate. 

Actinal intermediate area large; actinals not extending down arm. 
Actinal plates flat, polygonal; completely covered by short, conical 
spinules, about as high as wide. Some plates bearing short, excavate 
pedicellariae similar to those on inferomarginals. 



SMIIIISONIAK 
IN'SIITUTION 



MAY2 919F 



132 Proceedings of the Biological Society of Washington 




Fig. 2. Litonotaster intermedius (Perrier), from Oregon sta. 2574. 
Top, abactinal view; 1.1 X. Bottom, actinal view, 1.1 X. 



I 



The genus Litonotaster 



133 







Fig. 3. Litonotaster intermedius (Perrier), from Oregon sta. 2574. 
Top, abactinal view; 8.3 X- Bottom, actinal view; 7.9 X. 



134 Proceedings of the Biological Society of Washington 

Adambulacral plates large, rectangular (slightly wider than long), 
corresponding with adjacent actinals. Furrow margin straight, bearing 
six to eight short, blunt, compressed, subequal furrow spines. Three to 
four irregular rows of subambulacral spines, similar to those of actinals, 
covering rest of plate. Most plates bearing two or three valved excavate 
pedicellariae, ranging from as high as wide to almost three times as high 
as wide. 

Mouth plates large with long furrow margins. Each plate bearing 
twelve or thirteen furrow spines; first nine or ten similar to adambulacral 
furrow spines; next two slightly taller and more strongly compressed. 
Median spine about one and one-half times taller than other spines, very 
strongly compressed. Rest of plate covered by two to three irregular 
rows of spinules parallel to suture; spinules similar to those of actinals. 
Suture broad, prominent. 

Madreporite irregularly round, about twice as large as adjacent abac- 
tinals. 

Type: The type was originally deposited in the Museum of Compara- 
tive Zoology, Harvard Univ., but is no longer in their collection. It is 
presumed lost. 

Type-locality: 24°33'N,84°23'W, 3532 m, Blake sta. 31. 

Distribution: This species is known from the Gulf of Mexico (2379- 
3532 m) and from the Windward Passage between Cuba and Hispaniola 
(1958-3294 m). 

Discussion: The granulation of the abactinal plates is variable. In one 
specimen most of the plates are completely covered by granules. In 
another specimen, most of the plates have a very small, central naked 
area, bearing one to three scattered granules. In a third specimen, all 
the plates are surrounded by two or three rows of granules. The central 
areas are wide and completely naked. 

The superomarginals are usually in the vertical plane, but can be close 
to the horizontal plane or at any angle between. 

Litonotaster africanus new species 
Figures 4, 5 

Material studied: Holotype: R = 43 mm, r = 21 mm, R/r = 2; 01°09'- 
N, 07°45'E, 2525 m, R/V Pillsbury sta. 266, 17 May 1965, USNM 
E10863. 

Diagnosis: Abactinal plates covered by rounded, coarse, closely crowded 
granules. Small central area of some plates bare. Many plates bearing 
one or two moderately large, two or three valved excavate sugar-tong 
pedicellariae. Actinals covered by short, conical spinules, about as high 
as wide. Adambulacral plates bearing six to seven furrow spines. 

Description: Five arms. R = 43 mm; r =: 21 mm; R/r =z 2. 

General form stellate; arms strongly tapered; disk inflated; dorsal integ- 
ument thin, flexible; interbrachial arcs rounded. 

Abactinal plates small, flat, irregularly round, closely crowded; covered 



The genus Litonotaster 135 




Fig. 4. Litonotaster africanus, n. sp., holotype. Top, abactinal view; 
0.9 X. Bottom, actinal view, 0.9 X. 



136 Proceedings of the Biological Society of Washington 




Fig. 5. Litonotaster africanus, n. sp., holotype. Top, abactinal view; 
5.1 X. Bottom, actinal view, 6.1 X. 



The genus Litonotaster 137 

by rounded, coarse, closely crowded granules. Small central area of some 
plates bare. Many plates bearing one or two moderately large, two or 
three valved excavate sugar-tong pedicellariae. Interradial areas large. 
Papular areas small, inconspicuous; restricted to narrow bands running 
from bases of arms to center of disk; no papulae in center of disk. Six 
papular pores surrounding each plate, each pore containing a single 
papula. 

Twenty-two plates in each marginal series corresponding proximally, 
but tending to alternate distally. In interbrachial arcs lateral angle of 
each plate depressed; plates flattened. Plates becoming more angular 
distally. Superomarginal plates at an angle of about 30° from the horizon- 
tal. Superomarginals square; three or four pairs in contact medially. Each 
plate surrounded by one row of granules on three sides; two to four 
irregular rows on side adjacent to inferomarginals. Marginal granules 
rounded, similar to those on abactinals. Center of plates naked, except 
for scattered granules and excavate sugar-tong pedicellariae similar to 
those of abactinals. Inferomarginal plates square or pentagonal; covered 
by very short, conical spinules. Some plates bearing pedicellariae similar 
to those of abactinals. Terminal plate roundly triangular; distal end sur- 
mounted by short, blunt tubercle. 

Actinal intermediate area large; single row of actinals extending about 
half way down arm. Actinal plates small, flat, polygonal; completely 
covered by short, conical spinules, about as high as wide. Many plates 
bearing two or three valved pedicellariae similar to those found on 
abactinals. 

Adambulacral plates large, rectangular (slightly wider than long), 
corresponding with adjacent actinals. Furrow margin straight, bearing 
six or seven short, blunt, compressed, subequal furrow spines. Three to 
four irregular rows of subambulacral spines similar to those of actinals 
covering rest of plate. Most plates bearing two or three valved excavate 
sugar-tong pedicellariae, similar to but slightly larger than those found 
on other plates. 

Mouth plates large with long furrow margin. Each plate bearing 
thirteen furrow spines; first ten similar to adambulacral furrow spines; 
next two slightly taller and more strongly compressed. Median spine 
about one and one-half times taller than other spines, very strongly com- 
pressed. Rest of plate covered by two to three irregular rows of spinules 
parallel to suture; spinules similar to those of actinals. Suture broad, 
prominent. 

Madreporite round, about three times as large as adjacent abactinals. 

Type: United States National Museum, cat. no. E10863. 

Type-locality: Gulf of Guinea, off Sao Tome, 01°09'N, 07°45'E, 2525 
m, R/V Pillsbury sta. 266. 

Distribution: This species is known only from the type-locality. 

Discussion: Only a small number of the abactinal plates have a central, 
naked area, but no fewer than in some specimens of Litonotaster inter- 
medius, the type-species. 



138 Proceedings of the Biological Society of Washington 




Fig. 6. Litonotaster rotundigranulum, n. sp., holotype. Top, abactinal 
view; 0.7 X. Bottom, actinal view; 0.7 X- 



The genus Litonotaster 139 

This species is closely related to L. intermedius. It is distinguished from 
it by the coarse granules and the pedicellariae on the abactinal plates. 
Also, the holotype is considerably larger (R = 43 mm) than the largest 
specimen of L. intermedius known (R = 27 mm). 

Litonotaster tumidus H. L. Clark, 1920 
Figure 8 

Litonotaster tumidus H. L. Clark, 1920. pp. 83-85, pi. 3, figs. 3-6. — 
Madsen, 1951, p. 88. 

Material studied: Paratype: Peru, SW of Palominos Light House, 88 
miles, 5196 m. Albatross sta. 4672, MCZ 2717. 

Diagnosis: No abactinal pedicellariae. Actinal and inferomarginal 
plates covered by long spinules two to three times as high as wide. 
Adambulacral plates bearing four or five furrow spines. 

Type: The holotype is apparently lost. The extant paratypes are in 
the United States National Museum (cat. no. E705) and the Museum of 
Comparative Zoology (cat. nos. 2675 and 2717). 

Type-locality: Peru, SW of Palominos Light House, 88 miles, 5196 
m. Albatross sta. 4672. 

Distribution: This species is known from off the coast of Peru, 4066- 
5196 m. 

Discussion: Litonotaster tumidus is closely related to L. intermedius. 
It is distinguished from it by its tall actinal and inferomarginal spinules 
and the small number of adambulacral furrow spines (four or five). 

Litonotaster rotundigranulum new species 
Figures 6, 7 

Material studied: Holotype: R = 60 mm, r = 29 mm, R/r r= 2; 
23°54'N, 82°19'W, 1135-1184 m, R/V Gerda sta. G-448, 1 December 
1964, USNM E10862. 

Diagnosis: Disk large. Superomarginal plates prominent. Actinals cov- 
ered by large, hemispherical granules. Seven or eight furrow spines in 
proximal half of ambulacral furrow, five or six distally. No pedicellariae. 

Description: Five arms. R = 60 mm; r r= 29 mm; R/r = 2. 

General form stellate with large disk; arms tapering regularly to a 
bluntly pointed tip; disk slightly inflated; dorsal integument very thin, 
flexible; interbrachial arcs broadly curved. 

Small, flat, very thin abactinal plates irregularly round, closely crowded; 
covered by small, rounded granules. Small central area of most plates 
bare; some with one to four scattered granules. No secondary plates. 

Interradial areas large. Papular areas small, inconspicuous; restricted 
to narrow bands running from bases of arms to center of disk; no papulae 
in center of disk. Six papular pores surrounding each plate, each pore 
containing a single papula. 

Twenty-four to twenty-eight large superomarginal plates correspond- 
ing with inferomarginals in interbrachial arc, but irregular in shape, 



140 Proceedings of the Biological Society of Washington 



' --zM^--^- ■ ^10P^¥^A: 



|^^^^»:^^|-\ig>f 




W^wm^.- 





Fig. 7. Litonotaster rotundigranulum, n. sp., holotype. Top, abac- 
tinal view; 3.2 X. Bottom, actinal view; 3.0X. 



The genus Litonotaster 141 




im 

Fig. 8. Litonotaster tumidus H. L. Clark, from Albatross sta. 4672, 
actinal view; 9.0 X- 



number and correspondence in distal parts of arms; plates square in 
interbrachial arc, becoming wider than long distally; one to four pairs 
in contact medially. Lower half of each plate completely covered by 
large, hemispherical granules. Upper half bordered by a row of granules; 
center bare or with scattered granules. Lateral angle of superomarginals 
depressed so that each plate is flattened, moderately thin; becoming 
more angular distally. Superomarginals mainly in horizontal plane. Termi- 
nal plate moderately large, naked, swollen, heart-shaped; distal end 
notched, proximal end truncate. Twenty-six to thirty inferomarginal 
plates; always two more than in superomarginal series. Inferomarginals 
completely covered by large, hemispherical granules similar to those of 
superomarginals. 

Actinal intermediate area large. Plates large, rhombic except near 
marginals, where small, irregular. Actinals flat, well defined; completely 
covered by large, hemispherical granules. 

Adambulacral plates square or sometimes pentagonal, corresponding 
with adjacent actinals, which are twice as wide; furrow margin straight. 
Six or seven furrow spines in proximal half of ambulacral furrow, five 
or six distally; spines short, blunt, compressed, subequal. First row of 
actinal series of five or six very short, blunt spines; rest of actinal series 
consisting of three irregular rows of five or six large, hemispherical gran- 
ules like those of actinals. 

Mouth plates very large, with long furrow margin. Each plate with ten 
short, blunt furrow spines; first nine spines subequal, like adambulacral 



142 Proceedings of the Biological Society of Washington 



furrow spines; median spine only slightly larger. Rest of plate covered 
by large hemispherical granules like those of actinals except for short, 
blunt spines at suture; suture broad, prominent. 

Madreporite deeply furrowed, rhombic, about four times larger than 
surrounding plates; one-third the distance from center of disk to middle 
of interbrachial arc. 

No pedicellariae. 

Type: United States National Museum, cat. no. E 10862. 

Type-locality: Straits of Florida, on Pourtales Plateau, S. of Key West; 
23°54'N, 82°19'W, 1135-1184 m, R/V Gerda sta. G-448. 

Distribution: This species is known only from the type-locality. 

Discussion: This species is not closely related to the other known spe- 
cies of Litonotaster. It differs in its general body form, actinal granula- 
tion and lack of adambulacral pedicellariae. These differences are not 
sufficient to erect a new genus and Litonotaster can be expanded to 
accommodate this new species. 

Literature Cited 

Alcock, a. 1893. Natural history notes from H. M. Indian Survey 
Steamer "Investigator," Commander C. F. Oldham, R. N., 
commanding. Series 2, No. 7, An account of the collection 
of deep-sea Asteroidea. Ann. Mag. nat. Hist, (6) 11: 73-121, 
pis. 4-6. 

Clark, A. H. 1954. Echinoderms (other than holothurians ) of the 
Gulf of Mexico. Fish. Bull., 55: 373-379. 

Clark, H. L. 1920. Reports on an exploration off the west coasts of 
Mexico, Central and South America, and off the Galapagos 
Islands, ... by ... . "Albatross" during 1891 . . . 32. 
Asteroidea. Mem. Mus. comp. Zool. Harvard, 39 (3): 75- 
113, pis. 1-6. 

. 1941. Reports on the scientific results of the "Atlantis" 

Expeditions to the West Indies, under the joint auspices of 
the University of Havana and Harvard University. The echi- 
noderms (other than holothurians). Mem. Soc. cubana Hist, 
nat., 15 (1): 1-154, pis. 1-10. 

Fisher, W. K. 1911. Asteroidea of the North Pacific and adjacent 
waters. Part 1. Phanerozonia and Spinulosa. Bull. U. S. Nat. 
Mus., 76 ( 1 ) : 1-420, pis. 1-122. 

Madsen, F. J. 1951. Asteroidea. Rep. Swedish Deep-Sea Exp., 2(6): 
73-92, figs. 1-2. 

Perrier, E. 1884. Memoire sur les etoiles de mer recueillis dans la 
Mer des Antilles et la Golfe du Mexique durant les expedi- 
tions de dragage faites sous la direction de M. Alexandre 
Agassiz. Nouv. Arch. Mus. Hist. nat. Paris, (2) 6: 127-276, 
pis. 1-9. 

Verrill, a. E. 1899. Revision of certain genera and species of star- 
fishes. Trans. Conn. Acad. Arts Sci., 10: 145-234, pis. 24-30. 



BH 

1 

B4X 82, pp. 143-166 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



NOTES ON THE GONODACTYLUS SECTION OF THE 

FAMILY GONODACTYLIDAE (CRUSTACEA, 

STOMATOPODA), WITH DESCRIPTIONS OF FOUR 

NEW GENERA AND A NEW SPECIES 

By Raymond B. Manning 
Smithsonian Institution, Washington, D. C. 

The present report was stimulated by two separate studies, 
a review of the eastern Atlantic stomatopods and the prepara- 
tion of a description of a new gonodactylid found in the col- 
lections of the Division of Crustacea, Smithsonian Institution. 
Examination of the eastern Atlantic Protosquilla folini (A. 
Milne-Edwards) as part of a revision of the stomatopods from 
that area revealed that it differs from all others in the genus 
in two features: the articulated anterolateral plates of the ab- 
domen are absent and the submedian teeth of the telson lack 
denticles. These observations were noted in a report now in 
press on the stomatopods taken in the Gulf of Guinea in 1964 
and 1965 during two cruises of the R/V John Elliott Pilhbury 
( Manning, in press ) . 

Comparison of the new gonodactylid from Bougainville with 
other species formerly assigned to Protosquilla suggested that 
there were several previously unrecognized genera in that 
complex of species. This in turn led to this preliminary review 
of the genera allied to Gonodactylus. 

All of the genera recognized herein were included by Kemp 
( 1913 ) in the genus Gonodactylus, which, until very recently, 
included all species with a basally inflated, unarmed dactylus 
on the raptorial claw and the unusual ischiomeral articulation 
of the claw in which the merus projects posteriorly well beyond 
the articulation. Kemp recognized four separate groups in 
Gonodactylus and stated (p. 145): "... it seems that the 

9— Proc. Biol. Soc. Wash., Vol. 82, 1969 (143) 



^THSO/VT^ 

f JUN26t969) 



144 Proceedings of the Biological Society of Washington 

species here included in Gonodactylus fall into four natural 
groups of aproximately equal value, but these do not, I be- 
lieve, possess any greater claims to nomenclatorial distinction 
than do the groups in Lysiosquilla or in other genera." 

Kemp's four groups were as follows: 

Group I: This group contained the species now placed in 
Gonodactylus as restricted here. 

Group II : This assemblage comprised six species with highly 
modified telson and uropods. One of these species had been 
assigned to Mesacturus by Miers (1880), but this name had 
been overlooked by all subsequent workers until Holthuis 
(1967) resurrected it. The species of this Group are assigned 
herein either to Mesacturus or to a new genus, Gonodactylop- 
sis. 

Group III: This group included most of the species which 
Brooks (1886) had assigned to Protosquilla. Brooks had used 
the apparent fusion of the telson and the sixth abdominal so- 
mite as the basic character of the genus; as Kemp and others 
noted, however, this character did not prove valid. Four genera 
are recognized herein for these species — Protosquilla Brooks 
and three new genera. 

Group IV: Kemp assigned but one small, unique species to 
this group. Holthuis (1964) placed this species in Hoplos- 
quilla, and this assignment is recognized here. 

There are two basic telson types found among the genera 
treated; in all of the genera the telson is basically a flat plate 
ornamented with at least three rounded dorsal bosses, a median 
and two submedians. In the Gonodactylus-type telson, found 
in Gonodactylus, Gonodactylopsis, and Hoplosquilla, the telson 
and its dorsal carinae are inflated, and the posterior margin is 
produced into two or more pairs of usually widely-separated 
teeth (figure 1). In the Protosquilla-type telson, found in 
Chorisquilla, Echinosquilla, Haptosquilla, and Protosquilla, 
the marginal teeth, other than the submedians, are usually 
separated by short, narrow, incisions and the general outline 
is subquadrate (figure 6). Mesacturus, however, has two sep- 
arate telson types, one (figure 3) somewhat resembling the 
Gonodactylus-type, the other (figure 2) with a bifurcate me- 



GonodactyUd stomatopods 145 

dian posterior projection, possibly representing highly-modified 
submedian teeth, unique to the genus. 

Manning ( 1968 ) noted that the Family Gonodactylidae could 
be separated into two broad sections, one characterized by a 
Gonodactylus mien and the other by a resemblance to Pseu- 
dosquilla. The Gonodactylus section includes Gonodactylus, 
Hoplosquilla (the genera reported in more detail below), 
Odontodactylus, and Hemisquilla. 

Evolution in this section of the family apparently has cul- 
minated in five distinct groups of genera, all of which because 
of their similar morphology are believed to have arisen from 
a common ancestor. One line terminated in Hemisquilla, a 
genus which resembles Gonodactylus in basic facies, including 
basic telson shape, but which differs in lacking the subterminal 
ischiomeral articulation of the raptorial claw, in having a tri- 
angular rather than trilobed rostral plate, and in frequenting 
different types of habitats. Hemisquilla, with one species ex- 
tending northward to California, is basically a southern tem- 
perate genus, with populations in the south Atlantic, off Peru, 
Chile, and Australia; the Atlantic and Pacific species are dis- 
tinct. Also it is found on level bottoms, as opposed to the other 
genera in this section, all of which occur typically in rough, 
rocky areas or on coral reefs. 

Another line terminated in Odontodactylus, the only genus 
of this section with teeth on the raptorial claw. Manning 
(1967a) reviewed the species which occur in the Indo-West 
Pacific and western Atlantic regions. As in Gonodactylus and 
its allies, Odontodactylus usually occurs in rocky or coral reef 
habitats in tropical waters. The telson in Odontodactylus re- 
sembles that found in the Pseudosquilla section of the family. 

The third line includes three of the genera recognized 
herein, all of which share the Gonodactylus-type telson, Gono- 
dactylus, Gonodactylopsis, and Hoplosquilla. These three 
genera also have a subglobular cornea, rounded anterior ex- 
tensions of the lateral plates of the carapace, and subterminal 
articulation of the two segments of the uropodal exopod, with 
the proximal segment extending well beyond its articulation 
with the distal segment. Only Gonodactylopsis has a trispinous 



SMJfHSONtAN ^fi^^( 2 Q 10(^(2 



W 



146 Proceedings of the Biological Society of Washington 

rostral plate, similar to that found in Protosquilla and its allies; 
in Gonodactylus and Hoplosquilla the rostral plate usually is 
rounded anterolaterally. Of these genera, only Hoplosquilla 
lacks a mandibular palp. 

In Gonodactylus proper, which will not be treated further 
here, three groups of species can be distinguished, one con- 
sisting of those species related to G. chiragra, which possess 
three dorsal bosses on the telson, another of species of diminu- 
tive size, such as G. demanii, and a third in which there are five 
dorsal bosses on the telson, as in G. falcatus. Some of the small 
Indo-West Pacific species of Gonodactylus were treated by 
Manning ( 1967b ) , but the remainder of the Indo-West Pacific 
species are in need of revision. 

Mesacturus, to which only six species are assigned here, does 
not show a close relationship to the species of the Gonodactylus 
line or to the members of that line leading to Protosquilla and 
other genera. All of the species have telsons and uropods with 
unusual ornamentation and shape, as shown in figures 2 and 3. 
In most of the gonodactylids, the uropodal endopod normally 
is a straight, flattened paddle fringed with a single row of 
setae [exceptions are found in some species of Gonodactylus 
(see Manning, 1967b) and Gonodactylopsis, see below]. In 
species of Mesacturus the endopod is curved outward and the 
entire surface may be covered with setae. 

The fifth group of genera share the Protosquilla-type telson, 
concave or straight anterior margins of the lateral plates of the 
carapace, a sharply trispinous rostral plate, and a terminal 
articulation of the segments of the uropodal exopod. One genus, 
Haptosquilla, has retained the rounded cornea found in Gono- 
dactylus; it also differs from the other genera with the Protos- 
quilla-type telson in having the submedian teeth separated by 
a narrow fissure so that the anterior portions of the inner edges 
of the teeth are appressed for much of their length. In the other 
genera, Chorisquilla, Echinosquilla, and Protosquilla, the cor- 
nea is broadened or even bilobed and the submedian teeth of 
the telson are not appressed. Protosquilla, an offshoot of Choris- 
quilla, differs from all other genera in the family in having the 
articulated anterolateral plates of the telson suppressed; it 



Gonodactylid stomatopods 147 

also lacks submedian denticles, and the inner margins of the 
submedian teeth are lined with fine setae (fig. 4b). Echinos- 
quilla, which also seems to be an offshoot of the Chorisquilla 
line, is unique in having erect dorsal spines on the uropodal 
endopod, fleshy apices on the dorsal spines of the telson, and 
in having both the submedian and intermediate margins of the 
telson lined with fixed spines. 

Of these genera, excluding Odontodactylus and Hemisquilla, 
only Gonodactylus occurs in the Americas. The most common 
American species, G. oerstedii Hansen, is very similar to G. 
chiragra (Fabricius), the most common Indo-West Pacific 
species; oerstedii and all American species differ from chiragra 
and all Indo-West Pacific species of Gonodactylus in having 
an accessory carina on the inner surface of each intermediate 
carina on the telson. The American species all have arisen from 
a single species which must have been very close to chiragra. 

Only Protosquilla occurs in the tropical waters of the eastern 
Atlantic region, where it is extremely abundant in suitable 
habitats. It would seem that this genus has replaced or ex- 
cluded Gonodactylus from this area. 

The remainder of the genera occur only in the Indo-West 
Pacific region, but relatively few of the species involved have 
wide ranges. 

The key to genera of the Gonodactylus section given below 
will serve to distinguish all of the genera reported herein; 
Odontodactylus and Hemisquilla have been excluded from the 
key. 

In the keys given below, taxa not further treated are set 
apart in brackets; within the brackets I have included the cur- 
rent name of the genus or species, reference to that taxon in 
Kemp (1913), if applicable, other pertinent references, and 
notes on geographic distribution. Original references cited by 
Kemp (1913) are not repeated here. An asterisk preceding a 
specific name indicates that I have examined specimens of 
that species. 

Figures 4 and 8 were drawn by my wife Lilly, with the par- 
tial support of the Office of Oceanography and Limnology, 
Smithsonian Institution. The remainder of the illustrations are 
from the literature. 



148 Proceedings of the Biological Society of Washington 

I am indebted to Horton H. Hobbs, Jr., Smithsonian Institu- 
tion, and L. B. Holthuis, Rijksmuseum van Natuurlijke His- 
toric, Leiden, The Netherlands, for reading the manuscript; 
their comments materially improved the final draft. 

Key to Genera of Gonodactylus Section of Family 

GONODACTYLIDAE GlESBRECHT, 1910 

1. Anterior margins of lateral plates of carapace convex, extending 

well anterior to base of rostral plate ( fig. 1 ) 2 

Anterior margins of lateral plates of carapace straight or slightly 
concave, not extending anteriorly past base of rostral plate ( fig. 7 ) 5 

2. Mandibular palp absent; inner margin of endopod and distal seg- 
ment of exopod of uropod with sharp, fixed spines 

- [Hoplosquilla Holthuis, 1964; 

monotypic; Hoplosquilla acanthurus (Tattersall, 1906); Kemp, 
1913; Ceylon]. 

Mandibular palp present; inner margin of endopod and distal seg- 
ment of exopod of uropod with setae, not fixed spines 3 

3. Rostral plate not trispinous, anterolateral angles rounded or acute 

but not spiniform [Gonodactylus Berthold, 1827; 

about 26 species; pantropical except eastern Atlantic]. 

Rostral plate sharply trispinous 4 

4. Distalmost spines on outer margin of proximal segment of uropodal 
exopod enlarged, strongly recurved (fig. 3 ) ; propodus of claw lack- 
ing movable spine proximally on inner margin; apex of uropodal 

endopod curved outward Mesacturus, p. 150. 

Distalmost spines on outer margin of proximal segment of uropodal 
exopod not enlarged or strongly recurved ( fig. 1 ) ; propodus of 
claw with proximal movable spine on inner margin; apex of uro- 
podal endopod curved inward Gonodactylopsis, p. 149. 

5. Abdomen without articulated anterolateral plates (fig. 4a); inner 
edge of submedian teeth of telson lined with setae, not denticles 

(fig. 4b) Protosquilla, p. 153. 

Abdomen with articulated anterolateral plates (fig. 4c); inner 
edge of submedian teeth of telson lined with denticles or teeth ._.. 6 

6. Dorsal spines of telson with fleshy apices; uropodal endopod with 

erect dorsal spines (fig. 5) Echinosquilla, p. 155. 

Dorsal spines of telson, if present, lacking fleshy apices; uropodal 
endopod unarmed dorsally 7 

7. Cornea flattened; posterior margin of telson divided into 2 halves 
by V-shaped median emargination ( fig. 6 ) ; submedian denticles of 

telson long, slender Chorisquilla, p. 157. 

Cornea subglobular; posterior margin of telson divided into 2 
halves by narrow median fissure, with anterior edges appressed 

(fig. 7); submedian denticles of telson short, broad 

Haptosquilla, p. 159. 



Gonodactylid stomatopods 149 

Gonodactylopsis new genus 
Figure 1 

Diagnosis: Rostral plate sharply trispinous; cornea subglobular; ante- 
rior margins of lateral plates of carapace convex, extending anteriorly 
beyond base of rostral plate; ischiomeral articulation of claw not termi- 
nal; propodus of claw with proximal movable spine; dactylus of claw 
unarmed; articulation of propodus and dactylus inflated; mandibular 
palp present; articulated anterolateral plates of abdomen present; posterior 
margin of sixth abdominal somite straight or convex; telson of Gonodac- 
tylus-type, with 2 or 3 pairs of marginal teeth, submedians with move- 
able apices; submedian denticles present in 1 species, absent in other; 
proximal segment of uropodal exopod extending beyond articulation with 
distal segment, movable spines on outer margin of proximal segment of 
uropodal exopod straight or slightly curved; uropodal endopod curved 
inward, setation unusual; uropodal exopod and endopod without fixed 
spines on inner margin. 

Number of species: Two, distinguished in the key, below. 

Range: Indo-West Pacific region. 

Type-species: Gonodactylus herdmani Tattersall, 1906. 

Gender: Feminine. 

Etymology: The name is derived from the combination of Gonodac- 
tylus and -opsis, Greek, meaning like, referring to the resemblance of this 
genus to Gonodactylus. 

Remarks: Gonodactylopsis resembles both Gonodactylus and Mesacturus 
in having the anterior margins of the lateral plates of the carapace sloping 
forward beyond the base of the rostral plate and in having the proximal 
segment of the uropodal exopod extending beyond the articulation with 
the distal segment. It differs from Gonodactylus in having the rostral 
plate sharply trispinous; in none of the species of Gonodactylus does the 
shape of the rostral plate approach that of Gonodactylopsis. Gonodac- 
tylopsis lacks the enlarged, recurved spines on the outer margin of the 
uropodal exopod that are characteristic of Mesacturus. 

Gonodactylopsis also resembles Hoplosquilla, but differs in having a 
mandibular palp and in lacking fixed spines on the inner margins of the 
uropodal exopod and endopod. 

Gonodactylopsis can be distinguished from the genera with the Protos- 
quilla-tyTpe telson, each of which also has a trispinous rostral plate, by the 
subterminal articulation of the segments of the uropodal exopod. 

The uropodal endopod of Gonodactylopsis is curved inward, not out- 
ward as in Mesacturus, and the marginal setation is incomplete. 

Both of the species assigned to this genus are extremely rare. 

Key to species of Gonodactylopsis 
1. Submedian denticles of telson absent; dorsal surface of telson with 
3 longitudinal keels and a few rounded tubercles 

"* [Gonodactylopsis herdmani (Tattersall, 1906); Kemp, 1913; Cey- 
lon]. 



150 Proceedings of the Biological Society of Washington 




Fig. 1. Gonodactylopsis herdmani (Tattersall, 1906). a, sixth ab- 
dominal somite and telson; b, uropod; c, rostral plate (from Tattersall, 
1906). 



Submedian denticles of telson present; dorsal surface of telson with 
3 longitudinal keels and large, sharp spinules 

"^[Gonodactylopsis drepanophora (de Man, 1902); Kemp, 1913; 
Ternate, Molucca Islands; off Timor]. 

Mesacturus Miers, 1880 
Figures 2, 3 

Mesacturus Miers, 1880, p. 46.— Holthuis, 1967, p. 37. 

Diagnosis: Rostral plate sharply trispinous; cornea subglobular; an- 
terior margins of lateral plates of carapace convex, extending anteriorly 
beyond base of rostral plate; ischiomeral articulation of claw not termi- 
nal; propodus of claw lacking proximal movable spine; dactylus of claw 
unarmed; articulation of propodus and dactylus inflated; mandibular 
palp present, three-segmented; articulated anterolateral plates of abdo- 
men present; posterior margin of sixth abdominal somite straight; telson 
of unusual shape, ornamentation bizarre in some species; submedian 
marginal teeth of telson, if present, with movable apices; submedian 
denticles usually present; proximal segment of uropodal exopod extend- 
ing beyond articulation with distal segment, distalmost spines in outer 
margin of proximal segment enlarged, strongly recurved, uropodal endo- 
pod abnormal in shape and setation, usually curved outward, lacking 
dorsal spines; uropodal exopod and endopod lacking fixed spines on inner 
margin. 



Gonodactylid stomatopods 



151 




Fig. 2. Mesacturus furcicaudatus (Miers, 1880). a, last abdominal 
somite, telson, and uropod, dorsal view; b, same in oblique lateral view; 
c, telson, in posterior view (from Komai, 1940). 



Number of species: Six, distinguished in the key given below. 

Range: Indo-West Pacific region. 

Type-species: Gonodactylus furcicaudatus Miers, 1880, by monotypy. 

Remarks: Mesacturus can be distinguished from all other genera by the 
enlarged, strongly recurved distal spines on the outer margin of the 
proximal segment of the uropodal exopod. In addition, the unusual shape 
of the telson and the bizarre setation and shape of the uropods are help- 
ful in recognizing the members of the genus. 

Two general types of telson shape are found in the genus. In one 
(figure 2) the basal portion of the telson is broader than long and bears 
a slender, bifurcate posterior projection which arises from the middle of 
the posterior margin; the presence of a movable spine at the apex of each 
bifurcation suggests that these are modified submedian teeth. In the 
other type ( figure 3 ) the shape is reminiscent of that found in Gonodac- 
tylus, but the dorsal bosses are more numerous and often more elongate. 
The telson and uropods of members of this genus are highly specialized 
but their functions are unknown. 

Serene (1952) has shown that M. spinosocarinatus (Fukuda, 1910) 
has as its synonyms Gonodactylus strigatus Hansen, 1926, and G. demanii 
var. pruvotae Gravier, 1930. 

Key to species of Mesacturus 

1. Telson abnormal in shape, with narrow basal portion and slender, 

bifurcate median projection extending posteriorly (fig. 2) 2 

Telson normal in shape, length and width subequal, without bifur- 
cate process (fig. 3) 3 



152 Proceedings of the Biological Society of Washington 




Fig. 3. Mesacturus crinitus (Manning, 1962). a, last abdominal 
somite and telson; b, uropod, ventral view; c, uropod, dorsal view (from 
Manning, 1962). 



Gonodactylid stomatopods 153 

2. Posteromedian bifurcate process of telson slender, smooth, bifurcate 
for all of its length 

[Mesacturus furcicaudatus (Miers, 1880); Kemp, 1913; Komai, 
1940; Borodino Islands, south of Japan; Makatea, Polynesia; Banda 
Sea; off Lucipara Group, Indonesia]. 
Posteromedian bifurcate process of telson broad, spinous and heavily 

setose, with apical bifurcation only 

"^[Mesacturus kempi (Odhner, 1923); Ellice Islands]. 

3. Intermediate marginal teeth of telson small; lateral teeth present, 

small; dorsal surface of telson with 9 crowded longitudinal ridges 

"^[Mesacturus spinosocarinatus (Fukuda, 1910); Kemp, 1913; 
Serene, 1949, 1952; Japan; New Caledonia; near Saleyer, south of 
Celebes, Indonesia; Viet Nam; Queensland, Australia]. 
Intermediate marginal teeth of telson well-developed; lateral teeth 
absent; dorsal surface of telson not ornamented with 9 crowded 
longitudinal ridges 4 

4. Numerous intermediate denticles present on telson, 6-10 in 2 series 

on submedian tooth and 6-8 on intermediate tooth 

^[Mesacturus fimbriatus (Lenz, 1905); Kemp, 1913; Manning, 
1962; Zanzibar, Seychelles Islands]. 

Intermediate margin of submedian and intermediate teeth each 
with 1 denticle 5 

5. Uropodal endopod completely setose ventrally; proximal segment of 

uropodal exopod with dorsal patch of setae 

"^[Mesacturus brevisquamatus (Paul'son, 1875); Kemp, 1913; 
Manning, 1962; Red Sea]. 

Uropodal endopod lacking ventral setae; proximal segment of 

uropodal exopod without dorsal patch of setae 

^[Mesacturus crinitus (Manning, 1962); Seychelles Islands]. 

Protosquilla Brooks, 1886 
Figure 4a-b 

Diagnosis: Rostral plate sharply trispinous; cornea flattened; anterior 
margins of lateral plates of carapace straight or concave, not extending 
anteriorly beyond base of rostral plate; ischiomeral articulation of claw 
not terminal; propodus of claw lacking proximal movable spine; dactylus 
of claw unarmed; articulation of propodus and dactylus inflated; mandib- 
ular palp present, two-segmented; articulated anterolateral plates of 
abdomen absent (fig. 4a); posterior margin of sixth abdominal somite 
concave medially; posterior margin of telson divided into 2 halves by 
V-shaped median emargination; telson of Protosquilla-type, with 3 pairs 
of marginal teeth; telson lacking submedian denticles, inner margin of 
each submedian tooth lined with long, fine, fixed setae (fig. 4b); proxi- 
mal segment of uropodal exopod articulating terminally with distal seg- 
ment, movable spines on outer margin of proximal segment straight or 
slightly curved; uropodal endopod enlongate, with normal complement 
of setae, lacking fixed spines on inner margins. 



154 Proceedings of the Biological Society of Washington 






Fig. 4. Protosquilla folini (A. Milne-Edwards, 1867). a, first ab- 
dominal somite, lateral view; b, submedian teeth and denticles o£ tel- 
son, ventral view. Echinosquilla guerini (White, 1861), c, first abdominal 
somite, lateral view; d, submedian teeth and denticles of telson, ventral 



Number of species: One. 

Range: Eastern Atlantic Ocean, from the Cape Verde Islands to the 
Congo. 

Type-species: Gonodactylus folini A. Milne-Edwards, 1867, by subse- 
quent designation by Holthuis, 1967, p. 36. 

Remarks: Protosquilla differs from all other genera in the Gonodacty- 
lidae in having the articulated anterolateral plates of the abdomen totally 
suppressed (fig. 4a) and in having the submedian denticles of the telson 
replaced by long, thin setae (fig. 4b). It is the only representative of 
the Gonodactylus section of the family to occur in the eastern Atlantic 
Ocean where it seems to have completely replaced or excluded members 
of the genus Gonodactylus. 



Gonodactylid stomatopods 155 

Its striking similarity in features other than the two mentioned above 
to Chorisquilla trigibbosa (Hansen, 1926) suggests that it may have 
been derived from a similar species. 

Protosquilla elongata Brooks, 1886, based on a postlarva, is a synonym 
oi Protosquilla folini (A. Milne-Edwards, 1867). 

Echinosquilla new genus 
Figures 4c-d, 5 

Diagnosis: Rostral plate sharply trispinous; cornea flattened; anterior 
margins of lateral plates of carapace straight or concave, not extending 
anteriorly beyond base of rostral plate; ischiomeral articulation of claw 
not terminal; propodus of claw with proximal movable spine; dactylus 
of claw unarmed; articulation of propodus and dactylus of claw inflated; 
mandibular palp present, three-segmented; articulated anterolateral 
plates of abdomen very small; posterior margin of sixth abdominal somite 
concave medially; posterior distal margin of telson divided into 2 halves 
by V-shaped median emargination; telson of Protosquilla-type, with 2 
pairs of marginal teeth, submedians with movable apices; submedian 
denticles absent, inner margin of each submedian tooth lined with long, 
large, fixed spines; proximal segment of uropodal exopod articulating 
terminally with distal segment, movable spines on outer margin of proxi- 
mal segment straight or slightly curved; uropodal endopod elongate, 
with normal complement of setae, dorsal spines present; uropodal exopod 
and endopod lacking fixed spines on inner margin. 

Number of species: One. 

Range: Indo-West Pacific region, from scattered localities including 
Japan, Hawaii, Marquesas and Fiji Islands, in the Pacific Ocean, and 
Mauritius Island and Cosmoledo Atoll in the Indian Ocean. 

Type-species: Gonodactylus guerini White, 1861, by monotypy. 

Gender: Feminine. 

Etymology: The name is derived from the Latin, echinus, sea urchin, 
in combination with the generic name Squilla. 

Remarks: Echinosquilla can be distinguished from all other gonodac- 
tylids by the fleshy apices on the dorsal spines of the telson and the erect 
spines on the uropodal endopod. The wide submedian cleft of the telson 
as well as most of its general features suggest a close relationship with 
Chorisquilla; none of the species asigned to that genus have numerous 
long denticles lining the submedian and intermediate areas of the telson 
margin (figure 4d). 

The single species assigned to Echinosquilla, E. guerinii, is widely dis- 
tributed in the Indo-West Pacific region. It occurs in moderate depths, 
from less than 2 m to over 218 m. It has been suggested by Carie (1915) 
that the species uses the telson and uropods to block the entrance to its 
burrow; Carie pointed out the general resemblance of the telson to the 
echinoid Echinometra. 

Brooks' figure of E. guerini, published in 1886, is included here be- 



156 Proceedings of the Biological Society of Washington 




Fig. 5. EchinosquiUa merini (White, 1861). (from Brooks, 1886). 



Gonodactylid stomatopods 



157 




b 



Fig. 6. Chorisquilla lenzi (Holthuis, 1941). a, anterior portion of 
body; b, last two abdominal somites, telson, and uropod (from Kemp, 
1913). 



cause it portrays the basic facies of the species very well; unfortunately, 
the erect spines on the uropodal endopod are not clearly illustrated. 

Chorisquilla new genus 
Figure 6 

Diagnosis: Rostral plate usually sharply trispinous; cornea flattened; 
anterior margins of lateral plates of carapace straight or concave, not 
extending anteriorly beyond base of rostral plate; ischiomeral articula- 
tion of claw not terminal; propodus of claw with proximal movable 
spine; dactylus of claw unarmed; articulation of propodus and dactylus 
inflated; mandibular palp present; articulated anterolateral plates of 
abdomen very small; posterior margin of sixth abdominal somite concave 
medially; telson of Protosquilla-type, posterior margin divided into two 
halves by V-shaped median emargination; telson with 2 or 3 pairs of 



158 Proceedings of the Biological Society of Washington 

marginal teeth, submedians with movable apices; submedian denticles of 
telson long, slender, movable; proximal segment of uropodal exopod 
articulating terminally with distal segment, movable spines on outer 
margin of proximal segment straight or slightly curved; uropodal endopod 
elongate, with normal complement of setae, lacking dorsal spines; uro- 
podal exopod and endopod lacking fixed spines on inner margin. 

Number of species: Eight, distinguished in the key given below. 

Range: Indo-West Pacific region. 

Type-species: Gonodactylus excavatus Miers, 1880. 

Gender: Feminine. 

Etymology: The name is derived from the Greek, choris, meaning apart, 
in combination with the generic name Squilla. The name alludes to the 
separate submedian teeth of the telson. 

Remarks: Chorisquilla and the closely related Protosquilla and Echino- 
squilla share the flat, Protosqiiilla-type telson and numerous other features 
with Haptosquilla. Chorisquilla can be distinguished from Haptosquilla 
by the flattened or bilobed cornea, by the presence of the proximal 
movable spine on the propodus of the claw, and by the shape of the telson 
the posterior margin of which in Chorisquilla is divided into two halves 
by a V-shaped median emargination (fig. 6). Chorisquilla lacks the 
fleshy apices on the dorsal surface of the telson, the numerous long sub- 
median and intermediate denticles, and the erect spines on the uropodal 
endopod which are both characteristic of Echinosquilla. The articulated 
anterolateral plates of the abdomen will distinguish Chorisquilla from 
Protosquilla. 

Gonodactylus tweediei Serene, 1950, is here synonymized with G. 
trigibhosus Hansen, 1926; Hansen's species appears to be based on a 
juvenile and the the two major differences, relative shape of the rostral 
plate and the dorsal bosses of the telson, exhibited by the two nominal 
species can be attributed to differences in age. 

Although the submedian denticles of the telson appear to be movable 
in the species I have examined, C. spinosissima and C. trigibbosa, some 
of the illustrations in the literature seem to show that the denticles in 
other species may be fixed (vide Serene, 1947, pi. 4). 

Key to species of Chorisquilla 

1. Telson with 2 pairs of marginal teeth 2 

Telson with 3 pairs of marginal teeth . 



Dorsal surface of telson with numerous long spinules 

"^ [Chorisquilla spinosissima (Pfeffer, 1888); Kemp, 1913; Holthuis, 
1941; scattered localities between Japan, Australia, and the Red 
Sea]. 

Dorsal surface of telson not armed with numerous long spinules .— 
Dorsal surface of telson with numerous longitudinal ridges extend- 
ing to posterior margin 

* [Chorisquilla gyrosa ( Odhner, 1923 ) ; Gilbert Islands, Arno Atoll, 



Gonodactylid stomatopods 159 

Marshall Islands; Andaman Islands; Seychelles Islands; Diego 
Garcia, Indian Ocean]. 

Dorsal surface of telson not ornamented with numerous longitudinal 
ridges 4 

4. Submedian dorsal bosses of telson each extending to apex of sub- 
median teeth 

[Chorisquilla quinquelobata (Gordon, 1935); Christmas Island, 
Indian Ocean]. 

Submedian dorsal bosses of telson not extending to apices of sub- 
median teeth 5 

5. Submedian bosses of telson extending posteriorly beyond apex of 
median excavation in posterior margin; surface of all 3 dorsal bosses 

smooth; intermediate teeth of telson poorly-defined 

[Chorisquilla excavata (Miers, 1880); Bonin Islands; South China 
Sea; Indonesia; Viet Nam; Andaman Islands]. 

Submedian bosses of telson not extending posteriorly beyond apex 
of median excavation on posterior margin; surface of all 3 dorsal 

bosses tuberculate; intermediate teeth of telson well-defined 

[Chorisquilla tuberculata (Borradaile, 1907); Komai, 1938; Japan 
and Providence Island, Indian Ocean] . 

6. Dorsal surface of telson densely setose but not spinulose 

"^[Chorisquilla trigihhosa (Hansen, 1926); Serene, 1952 (as Gono- 
dactylus tweediei); Saleyer, Indonesia; Queensland, Australia; Lord 
Howe Island]. 

Dorsal surface of telson with numerous spinules 7 

7. Telson with pyriform median and submedian bosses; dorsal spinules 

straight 

[Chorisquilla brooksii (de Man, 1887); Kemp, 1913; Serene, 1947; 
Tidore and Tenimber Islands, Indonesia; South China Sea; Viet 
Nam] . 

Telson with subcircular median and submedian bosses; dorsal 

spinules recurved posteriorly 

[Chorisquilla hystrix (NobiH, 1899); Kemp, 1913; Beagle Bay, 
British New Guinea]. 

Haptosquilla new genus 
Figure 7 

Diagnosis: Rostral plate trispinous; cornea subglobular; anterior 
margins of lateral plates of carapace straight or concave, not extending 
anteriorly beyond base of rostral plate; ischiomeral articulation of claw 
not terminal; propodus of claw usually lacking proximal movable spine; 
dactylus of claw unarmed; articulation of propodus and dactylus of claw 
inflated; mandibular palp usually present; articulated anterolateral plates 
of abdomen well-developed; posterior margin of sixth abdominal somite 
concave medially; telson of Protosquilla-type, divided into 2 halves by 
narrow median fissure, with anterior edges appressed; telson with 3 or 4 



160 Proceedings of the Biological Society of Washington 




r '^1 







Fig. 7. Haptosquilla pulchella (Miers, 1880). a, anterior portion 
of body; b, last two abdominal somites, telson, and uropod (from Kemp, 
1913). 



pairs of marginal teeth, submedians with movable apices; submedian den- 
ticles short, broad, movable; proximal segment of uropodal exopod articu- 
lating terminally with distal segment, movable spines on outer margin of 
proximal segment straight or slightly curved; uropodal endopod elon- 
gate, with normal complement of setae, lacking dorsal spines; uropodal 
exopod and endopod lacking fixed spines on inner margins. 

Number of species: Thirteen, including one new species described 
herein; they may be distinguished by means of the key given below. 

Range: Indo-West Pacific region. 

Type-species: Gonodactylus pulchellus Miers, 1880. 

Gender: Feminine. 

Etymology: The name is from the Greek, hapto, to touch, in combina- 
tion with the generic name Squilla. The name alludes to the appressed 
submedian teeth of the telson. 

Remarks: The appressed submedian lobes of the telson will distinguish 
Haptosquilla from other members of the Gonodactylidae. The rounded 
cornea in the species of Haptosquilla will also serve to distinguish them 
from members of Chorisquilla, Echinosquilla, and Protosquilla, in which 
the cornea is either broadened or bilobed. 

The rostral plate of most species of Haptosquilla is sharply trispinous. 
However, in H. nefanda (Kemp) the anterolateral spines are not well 
developed; they are sharp and acute but not spiniform. 

Haptosquilla glyptocercus (Wood-Mason, 1875) has as its synonym 
Protosquilla cerebralis Brooks, 1886. H. lenzi (Holthuis, 1941) was pro- 
posed as a replacement name for the preoccupied Gonodactylus glaber 
Kemp, 1913, itself an emendation of Protosquilla glabra Lenz, 1905. 



Gonodactylid stomatopods 161 

Haptosquilla hamifera (Odhner, 1923) and H. pulchra (Hansen, 1926) 
may prove to be synonyms upon further study; pulchra is almost certainly 
based upon a juvenile. 

Key to species of Haptosquilla 

1. Telson with 3 pairs of marginal teeth 2 

Telson with 4 pairs of marginal teeth 4 

2. Median portions of fifth and sixth abdominal somites wrinkled; 

dorsal submedian bosses of telson extending to posterior margin 

"^ [Haptosquilla glyptocercus (Wood-Mason, 1875); Brooks, 1886; 
Kemp, 1913; Scattered localities between Japan and Australia; 
Andaman Islands]. 

Median portion of fifth abdominal somite smooth; dorsal sub- 
median bosses of telson not extending to posterior margin 3 

3. Dorsal surface of telson smooth between bosses 

"* [Haptosquilla lenzi (Holthuis, 1941); Kemp, 1913; Philippine 
Islands, Viet Nam, to the Indian Ocean and the Red Sea]. 
Dorsal surface of telson with well-marked depressions between 

bosses 

[Haptosquilla tanensis (Fukuda, 1911); Kemp, 1913; Japan]. 

4. Telson with prominent median boss only on dorsal surface, sub- 
median bosses replaced by short, rounded ridges . 



Telson with prominent median and submedian bosses on dorsal 
surface 6 

Dorsal ornamentation of telson not extending to posterior margin, 

posterior third of dorsal surface smooth 

[Haptosquilla pulchra (Hansen, 1926); Viet Nam; Borneo Bank]. 

Dorsal ornamentation of telson extending to posterior margin 

[Haptosquilla hamifera (Odhner, 1923); Fiji Islands]. 

Submedian bosses of telson extending to posterior margin 

[Haptosquilla ectypa (Miiller, 1887); Kemp, 1913; Bonin Islands; 

Ceylon; Mauritius]. 

Submedian bosses of telson short, not extending to posterior margin 

7 

Median portion of fifth abdominal somite corrugated 8 

Median portion of fifth abdominal somite smooth 10 

Submedian bosses of telson extending beyond mid-length of telson 

but not reaching posterior margin 

"^[Haptosquilla stoliurus (Miiller, 1887); Kemp, 1913; Western 
Australia and Malay Archipelago to Philippine Islands]. 
Submedian bosses of telson not extending posteriorly beyond mid- 
length of telson 9 

Rostral plate sharply trispinous; submedian bosses of telson oval 

at base - 

[Haptosquilla trispinosa (Dana, 1852); Kemp, 1913; South Pa- 
cific; Fiji Islands, Loyalty Islands, Australasian waters, and Viet 
Nam]. 



162 Proceedings of the Biological Society of Washington 

Anterolateral angles of rostral plate acute but not sharply spinous; 

submedian bosses of telson circular at base 

"^ [Haptosquilla tuherosa (Pocock, 1893); Kemp, 1913; Maccles- 
field Bank, South China Sea; Viet Nam] . 

10. Mandibular palp absent 

[Haptosquilla proxima (Kemp, 1915); Philippine Islands]. 
Mandibular palp present, two-segmented 11 

11. Ocular scales produced laterally, extending nearly to lateral rostral 
spine; anterior margins of carapace concave, anterolateral angles 

subacute 

"^ [Haptosquilla pulchella (Miers, 1880); Kemp, 1913; Philippine 
Islands and Australia to western Indian Ocean]. 

Ocular scales small, squarish, not produced laterally; anterior 
margins of carapace straight, anterolateral angles rectangular 12 

12. Anterolateral angles of rostral plate acute, blunt; dorsal bosses of 

telson smooth, non-setose 

^[Haptosquilla nefanda (Kemp, 1911); Kemp, 1913; Philippine 
Islands; Indonesia; Nicobar Islands; Andaman Islands; and 
Burma] . 

Anterolateral angles of rostral plate acute, sharp; dorsal bosses of 

telson with short, scattered setae 

"^Haptosquilla setifera new species. 

Haptosquilla setifera new species 
Figure 8 

Holotype: 1 $ , 14.0 mm; Bougainville, Solomon Islands; 1945; SI Div. 
Crustacea 125346. 

Paratype: 1 $ , 14.4 mm; data as in holotype; SI Div. Crustacea 125347. 

Description: Ocular scales small, subquadrate, lateral angles not 
markedly produced outward. 

Rostral plate with median spine slenderer and longer than laterals; 
lateral spines acute, sharp, short. 

Anterior margins of lateral plates of carapace straight, anterolateral 
angles rectangular, not produced anteriorly. 

Mandibular palp present, two-segmented. 

Outer margin of dactylus of claw notched proximally; propodus of 
claw with slender movable spine proximally. 

First 4 abdominal somites smooth, ornamented with at most an in- 
verted L-shaped groove anterolaterally; fifth somite with 3 pairs of 



Fig. 8. Haptosquilla setifera new species, holotype. a, anterior 
portion of body; b, last two abdominal somites, telson, and uropod ( setae 
omitted ) . 



Gonodactylid stomatopods 



163 




164 Proceedings of the Biological Society of Washington 

longitudinal swellings, outermost sharpest, median area smooth; sixth 
somite with 3 pairs of inflated carinae, each with posterior spine; surface 
of carinae irregular, wrinkled and pitted, several short setae present. 

Telson broader than long, of Protosquilla-type, with 3 oval dorsal 
bosses, not widely separate, submedians short, extending posteriorly about 
to apex of median cleft; dorsal bosses and surface of telson with broad, 
shallow pits, bosses irregular in outline; few short, fine setae present on 
bosses, dorsal surface of telson, and telson margin setae not so thick so as 
to obscure surface ornamentation; 4 pairs of blunt marginal teeth present, 
submedians with movable apices; numerous short, movable submedian 
denticles present, and 1 fixed denticle present between each of remaining 
marginal teeth. 

Uropod with 9 slender spines on outer margin of proximal segment of 
exopod, distalmost extending almost to end of distal segment; inner spine 
of basal prolongation smaller and shorter than outer. 

Color: Largely faded in both specimens, although the female shows 
traces of small black chromatophores arranged in a transverse band on 
each somite. 

Measurements: Male holotype, TL 14.0 mm, carapace length 2.5 mm, 
fifth abdominal somite width 2.5 mm, telson length 2.1 mm, width 2.5 
mm. Female paratype, TL 14.4 mm, carapace length 2.6 mm, fifth ab- 
dominal somite width 2.6 mm, telson length 2.3 mm, width 2.6 mm. 

Discussion: Both of the specimens upon which this description is based 
are obviously juveniles but the distinguishing features which they ex- 
hibit are not among those which might be expected to change with age. 
Even at this size this new species is distinguishable from all other species 
now in the genus. 

Haptosquilla setifera is closely related to H. nefanda (Kemp), H. 
pulchella (Miers), and H. proxima (Kemp). The presence of short setae 
on the dorsal bosses of the telson and the sharp anterolateral spines of 
the rostral plate will separate setifera from nefanda. The new species 
differs from pulchella in having many fewer setae on the telson, irregu- 
larly oval pitted bosses on the telson (in pulchella the bosses are smooth 
and round), truncated ocular scales, and rectangular anterolateral angles 
on the carapace; H. setifera is readily distinguishable from even the 
smallest specimen of pulchella (TL 19 mm) available for comparison. 
The mandibular palp will separate setifera from proxima, in which the 
palp has been suppressed. 

None of the other specimens of several species of Haptosquilla which 
I have examined have a proximal movable spine on the upper margin of 
the propodus of the claw. It may be characteristic of juveniles in this 
genus. 

Distribution: Known only from the type-locality, Bougainville, Solo- 
mon Islands. 

Etymology: The name is derived from the Latin, seta, referring to the 
short hairs present on the dorsal surface of the telson. 



Gonodactylid stomatopods 



165 



Literature Cited 

BoRRADAiLE, L. A. 1907. Stomatopoda from the western Indian Ocean. 
In The Percy Sladen Trust Expedition to the Indian Ocean 
in 1905, under the leadership of Mr. J. Stanley Gardiner. 
Trans. Linn. Soc. London, (2) 12: 209-216, pi. 22. 

Brooks, W. K. 1886. Report on the Stomatopoda collected by H.M.S. 
"Challenger" during the years 1873-1876. Rep. Voy. Chal- 
lenger, Zool., 16: 1-116, pis. 1-16. 

Carie, p. 1915. Note sur Gonodactylus (Protosquilla) Guerini White. 
Bull. Mus. Hist. Nat. Paris, 21: 151. 

Gordon, I. 1935. On two new species of Crustacea from Christmas 
Island. Ann. Mag. Nat. Hist., (10) 16: 629-637, figs. 1-3. 

Gravier, Ch. 1930. Sur une collection de crustaces ( Stomatopodes ) 
recueillis par Mme. Pruvot sur les cotes de la Nouvelle- 
Caledonie. Bull. Mus. Hist. Nat. Paris, (2) 2 (2): 214-216, 
fig. 1. 

Hansen, H. J. 1926. The Stomatopoda of the Siboga Expedition. 
Siboga Exped., Monogr. 35: 1-48, pis. 1-2. 

HoLTHUis, L. B. 1941. The Stomatopoda of the Snellius Expedition. 
Biological results of the "Snellius" Expedition, no. 12. Tem- 
minckia, 6: 241-294, figs. 1-9. 

— . 1964. Preliminary note on two new genera of Stomatopoda. 

Crustaceana, 7 (2): 140-141. 

. 1967. The stomatopod Crustacea collected by the 1962 and 

1965 Israel South Red Sea Expeditions. The Second Israel 
South Red Sea Expedition, 1965, Report No. 1. Israel Journ. 
Zool., 16: 1-45, figs. 1-7. 

Kemp, S. 1913. An account of the Crustacea Stomatopoda of the 
Indo-Pacific region, based on the collection in the Indian 
Museum. Mem. Indian Mus., 4: 1-217, 10 text figs., pis. 1-10. 

. 1915. On a collection of stomatopod Crustacea from the 

Philippine Islands. Philippine J. Sci., (D) 10 (3): 169-187, 
pi. 1. 

KoMAi, T. 1938. Stomatopoda occurring in the vicinity of Kii Penin- 
sula. Annot. Zool. Japon., 17 (3-4): 264-275, figs. 1-3. 

. 1940. Gonodactylus furcicaudatus Miers, a remarkable 

stomatopod. Annot. Zool. Japon., 19 (1): 47-50, fig. 1. 

Manning, Raymond B. 1962. Stomatopod Crustacea collected by the 
Yale Seychelles Expedition, 1957-1958. Postilla, no. 68: 1-15, 
figs. 1-2. 

. 1967a. Review of the genus Odontodactylus (Crustacea: 

Stomatopoda). Proc. U.S. Nat. Mus., 123 (3606): 1-35, figs. 
1-8, pi. 1. 

. 1967b. Notes on the demanii section of genus Gonodac- 
tylus Berthold with descriptions of three new species. Proc. 
U.S. Nat. Mus., 123 (3618): 1-27, figs. 1-8. 



166 



Proceedings of the Biological Society of Washington 



1968. A revision of the family Squillidae (Crustacea, 
Stomatopoda ) , with the description of eight new genera. Bull. 
Mar. Sci., 18 (1): 105-142, figs. 1-10. 

. In press. The stomatopod Crustacea. The R/V Pillsbury 

deep-sea Biological Expedition to the Gulf of Guinea, 1964- 
65. Stud. Tropical Oceanogr. 

MiERS, E. J. 1880. On the SquiUidae. Ann. Mag. Nat. Hist., (5) 5: 
1-30, 108-127, pis. 1-3. 

Odhner, T. 1923. Indopazifische Stomatopoden. Goteborg Vetensk. 
Samh. Handl., (4) 27 (4): 1-16, figs. 1-10. 

Serene, R. 1947. Sur les stomatopodes rare trouves en Indochina et 
n'existant pas dans les collections du Museum. Bull. Mus. 
Hist. Nat. Paris, 19 (5): 381-389, figs. 1-2, pis. 1-4. 

. 1949. Observations sur le Gonodactylus strigatus Hansen 

( crustace stomatopode ) . Bull. Soc. Zool. France, 74 ( 4-5 ) : 
225-231, 2 figs. 

. 1950. Deux nouvelles especes Indo Pacifiques de stomato- 
podes. Bull. Mus. Hist. Nat. Paris, (2) 22 (5): 571-572. 

. 1952. Etude d'une collection de stomatopodes de I'Aus- 

tralian Museum de Sydney. Rec. Australian Mus., 23 ( 1 ) : 
1-24, figs. 1-33, pis. 1-3. 

Tattersall, W. M. 1906. On the Leptostraca, Schizopoda and Sto- 
matopoda collected by Professor Herdman, at Ceylon, in 
1902. Supplementary Report XXXIII. In Herdman, Report 
to the Government of Ceylon on the Pearl Oyster Fisheries 
in the Gulf of Manaar, 5: 157-188, pis. 1-3. 



QH 

1 

B4X 

MH • 82, pp. 167-170 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



A NEW GENUS AND TWO NEW SPECIES 
OF ENTOCYTHERID OSTRACODS 
FROM ALABAMA AND MISSISSIPPI 

By H. H. Hobbs III 
Department of Zoology, Mississippi State University, State 
College 39762 and Gulf Coast Research Laboratory, Ocean 

Springs, Mississippi 

Described below are a new genus and two new species of 
ostracods epizootic on the crayfish Camharus diogenes dio- 
genes Girard. The members of the new genus have no close 
relationships with any others of the family, but, as in the two 
species of the genus Plectocy there, Hobbs III, 1965, each peni- 
ferum of the male supports a very long, slender penis which 
reaches the ventral extremity of the peniferum. Furthermore, 
the new genus is unusual in that it is the only one in which the 
spermatic loop rests horizontally in the peniferum. The clasp- 
ing apparatus resembles that of no other entocytherid. 

The other ostracod is sl member of the genus Ornithocy- 
there and is very similar to O. waltonae, Hobbs, Jr., 1967, the 
only previously known member of the genus in that the ventral 
portion of the peniferum is highly sclerotized and resembles the 
head and neck of a bird. The genus has a discontinuous dis- 
tribution—one portion of the range extends along the coastal 
plains of Virginia and Maryland, and the other, much farther 
inland, well above the fall line, in northern Mississippi. 

I would like to thank Dr. J. F. Fitzpatrick, Jr., who made the 
crayfish identifications and Dr. Horton H. Hobbs, Jr., for his 
assistance. Supported in part by NSF Grant GB-4719'to J. F. 
Fitzpatrick, Jr., and NSF Grant GB-7585 to Gulf Coast Re- 
search Laboratory. 

Saurocythere new genus 
Etymology: Saurocythere — Sslutos (Gr.), lizard; plus the generic name 
10— Proc. Biol. Soc. Wash., Vol. 82, 1969 (167) 

€4S0/V7^ 
26 1969 
7\RIE3, 



168 Proceedings of the Biological Society of Washington 

Cythere, referring to the resemblance of the peniferum of the male to 
the head of a Hzard. 

Diagnosis: Terminal tooth of mandible with cusps; no finger guard 
on copulatory complex of male; peniferum slender and elongate without 
accessory groove and extending ventrally beyond clasping apparatus; 
penis consisting of separate spermatic and prostatic elements, each one- 
fourth length of peniferum and reaching ventral extremity of peni- 
ferum; penis complex three times longer than least diameter of peni- 
ferum; spermatic loop situated horizontally and supported by heavily 
sclerotized bar; clasping apparatus not clearly divisible into vertical and 
horizontal rami, with external border sinous, bearing small excrescence 
at base of distal fourth; internal border entire; apical portion of clasping 
apparatus sHghtly expanded and serrate (fan-hke). 

Type-species: Saurocythere rhipis new species 

Saurocythere rhipis new species 
(Figs. 1, 4, 5) 

Etymology: rhipis (Gr.), fan, referring to the expanded fan-Uke distal 
portion of the clasping apparatus. 

Male: Eye pigmented, located approximately one-third shell length 
from anterior margin. Shell (Fig. 1) ovate with ventral margin entire, 
greatest height shghtly posterior to midlength. Length 0.442 mm., height 
0.294 mm. Setae scarce, occurring irregularly but singly near anterior and 
posterior margins. Mandible with distal row of five multicuspid teeth, 
proximal tooth with two cusps, distal with five, others with three. 

Copulatory complex (Fig. 5): Described in generic diagnosis; in addi- 
tion, dorsal finger long and slender, nearly one-half length of ventral 
finger and bearing two terminal spines, ventral finger slender and paral- 
lel to clasping apparatus ending in single terminal. 

Female: Unknown. 

Type-locality: Crayfish burrow in drainage ditch along State Route 14, 
0.6 mi. SE of Alabama-Mississippi state line, Pickens County, Alabama. 

Host: Cambarus d. diogenes Girard. 

Disposition of types: the holotypic male and a male paratype, the only 
known specimens are deposited in the United States National Museum, 
nos. 123880 and 123881, respectively. 

Range: The species is known only from the type-locaUty. 

Associates: An unidentified member of the genus Anklocythere. 

Ostracod Relationships: Saurocythere rhipis seems to be most closely 
alhed to the members of the genus Tlectocythere, Hobbs III, 1965. The 
principal resemblances are associated with the penis complex, which is 
composed of extremely long contiguous spermatic and prostatic elements 
in members of both genera. Although, the peniferum of Okriocythere 
cheia. Hart, 1964, exhibits a strong superficial resemblance to that of 
S. rhipis, the penis is much shorter, the spermatic loop is vertical, and 
the form of the clasping apparatus in both genera is unique. The hori- 



1 



Entocytherid ostracods 



169 




, .03mm , 



Figures 1-5. Figs. 1, 4, 5. Saurocythere rhipis new species: 1. Shell of 
holotypic male; 4. Ventral third of peniferiun of holotypic male; 5. 
Copulatory complex of holotypic male. Figs. 2, 3. Ornithocythere 
gypodes new species: 2. Shell of holotypic male; 3. Copulatory complex 
of holotypic male. 



zontally situated spermatic loop has not been observed to occur in any 
other entocytherid except S. rhipis. 

Ornithocythere gypodes new species 
(Figs. 2, 3) 
Etymology: gypodes (Gr.), vulture; so named because of resemblance 
of ventral portion of peniferum to the head of a vulture. 

Male: Eye pigmented, located approximately one-fourth shell length 
from anterior margin. Shell (Fig. 2) subelliptical, highest anterior to 
midlength with dorsal margin tapering to small eminence at posterior 
end of shell; ventral margin entire. Length 0.495, height 0.245. Terminal 
tooth of mandible with cusps. 



INSIITUTIOK 



MAY 2 9 1959 



170 Proceedings of the Biological Society of Washington 

Copulatory complex (Fig. 3) without finger guard; penifenim ex- 
tending ventrally beyond clasping apparatus with ventral portion highly 
sclerotized and produced into beak-like prominence directed anteriorly; 
accessory groove absent; penis complex with spermatic and prostatic 
elements propinquant throughout their length, not separated; penis com- 
plex slightly longer than anterior-posterior dimension of peniferum at 
level of base of penis. Two tubercles located on proximo ventral angle of 
beak-like prominence, a third one on ventral margin near apex; clasping 
apparatus not divisible into vertical and horizontal rami, with proximal 
and distal portions disposed at angle of approximately forty degrees; in- 
ternal and external borders entire except distally, there four annulations 
immediately preceding three apical denticles. Dorsal finger moderately 
slender, bearing one terminal spine; ventral finger slender and subparallel 
to clasping apparatus, bearing one terminal spine. 

Female: Unknown. 

Type-locality: Burrow in flood plain of Tombigbee River, one-fourth 
mi. east of river, 1 mi. west Alabama-Mississippi state line, Lowndes 
County, Mississippi. 

Host: cambarus d. diogenes Girard. 

Disposition of types: The dissected male holotype is deposited in the 
United States National Museum, no. 123879. 

Range: The species is known only from the type locaUty. Repeated 
efforts to obtain additional specimens of this ostracod at the type-locality 
and other localities along the Tombigbee River have, until now, been 
futile. 

Relationships: Quite clearly, Ornithocythere gypodes has its closest 
affinities with O. waltonae, Hobbs, Jr., 1967. The penifera are distinctly 
similar in that both emphatically resemble the head of a bird. The penis 
complexes are markedly similar, but the penis of O. gypodes is longer 
than that of O. waltonae. Likewise, O. gypodes is similar to Okriocythere 
cheia, Hart, 1964, to the members of the genus Geocythere, and less so 
to the members of the genera Ascetocythere, Hart, 1962, and Plectocy- 
ihere, Hobbs III, 1965. The combination of the more ventrally situated 
penis, the three spines located on the beak-like prominence of the peni- 
ferum, and the quite distinct clasping apparatus are unique to O. gypodes. 

Literature Cited 

Hart, C. W., Jr. 1962. A revision of the ostracods of the family 
Entocytheridae. Proc. Acad. Nat. Sci. Phila., 114: 121-147. 

. 1964. Two new entocytherid ostracods from the vicinity 

of Washington, D. C. Proc. Biol. Soc. Wash., 77: 243-246. 

Hobbs, Horton H., Jr. 1967. A new genus and three new species of 
ostracods with a key to genus Dactylocy there (Ostracoda: 
Entocytheridae). Proc. U. S. Nat. Mus., 122 (3587): 1-10. 

Hobbs, H. H., III. 1965. Two new genera and species of the ostracod 
family Entocytheridae, with a key to the genera. Proc. Biol. 
Soc. Wash., 78: 159-164. 




QH 

i 

MH I. 82, pp. 171-176 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



THE MONOGENEAN PARASITES OF AFRICAN FISHES. 

IX. A NEW GENUS, GUSSEVSTREMA, RECOVERED 

FROM THE GILLS OF TERAPON JARBUA (FORSKAL) 

FROM SOUTH AFRICA^ 

By Charles E. Price and Eric S. McClellan 

Department of Biology, Millersville State College, 

Millersville, Pennsylvania 

The monogenetic trematodes of African freshwater fishes have 
received considerable attention in recent years. In 1961 only 
four species of Monogenea had been described. At present 
approximately 60 species are known from the continent. This 
study is concerned with a new genus, Gussevstrema, recovered 
from the gills of Terapon jarbua (Forskal) from the Republic 
of South Africa. 

Host specimens were first frozen and then preserved in 3 per- 
cent formahn prior to shipment to the United States. Branchial 
materials and recovered parasites were then treated as pre- 
scribed by Price ( 1966) and measurements made as outlined by 
Price and McMahon (1967). Appropriate measiurements and 
illustrations were made with the aid of a calibrated filar microm- 
eter ocular and a camera lucida, respectively. Anatomical terms 
employed are those recommended by Hargis (1958) and by 
Price and Arai (1967). Average measurements are given first, 
followed by minimum and maximum values enclosed in paren- 
theses. All measurements are expressed in microns. 

Gussevstrema new genus 

Dactylogyridae, Ancyrocephalinae: Forms of moderate size, provided 
with a smooth cuticle. One pair of eyespots; comprising granules in 
anterior cephalic region. Cephalic lobes vestigial to lacking. Head organs 



1 This research supported by: (1) the Natal Parks, Game and Fish preserva- 
tion Board, Pietermaritzburg, Republic of South Africa, (2) Millersville State 
College and (3) a research grant from the American Philosophical Society (# 4956 
— Penrose fimd). 

11— Proc. Biol. Soc. Wash., Vol. 82, 1969 (171) 



f JUN26 1969) 



172 Proceedings of the Biological Society of Washington 

( on either side ) consist of four glandular structures connected to a larger 
pharyngeal gland. Haptor provided with two pairs of anchors. Bases of 
each anchor pair connected by a transverse haptoral bar, the bars con- 
nected to each other by multiple articulations; dorsal bar composed of 
three portions fused sohdly together. Hooks 14 ( seven pairs ) . Copulatory 
complex composed of a cirrus and a basally articulated accessory piece. 
Testis large, postovarian. Vagina and seminal receptacle not observed 
with certainty. Vitellaria exceptionally dense, composed of small granules. 
Intestinal hmbs apparently confluent posteriorly. 

Gussevstrema amacleithrium new genus, new species 

Host: Terapon jarhua (Forskal). 

Locality: Various bodies of water near Natal, South Africa. 

Location on host: Gill filaments. 

Number of specimens studied: Three. 

Holotype: USNM Hehn. CoU. No. 71359. 

Paratypes: In collection of senior author. 

Description: A small dactylogyrid provided with a thin, smooth cuticle, 
length 385 (377-398): greatest width of body 124 (118-129). One pair 
of eyespots, each provided with a lens. A few eyespot granules scattered 
in anterior cephaHc region. Cephahc lobes essentially lacking. Head 
organs ( on either side ) consist of four glandular structures all connected 
together by a fine duct; the duct continues posteriorly to terminate in 
a larger pharyngeal gland. Pharynx prominent, subspherical in dorsal 
view. Peduncle short and stout, setting the haptor well off from the 
body proper; haptor bilobed, with a small cleft in center of posterior 
border. Haptor apparently somewhat denser in central region, providing 
a differentiated site for attachment of anchors and bars. (Fig. 1, whole 
mount). Haptor provided with two pairs of anchors, one pair dorsal, the 
other ventral in position (Figs. 2, 3). Ventral anchors larger than dorsals; 
each ventral anchor composed of: (1) a soHd base provided with vestig- 
ial roots, (2) a sohd point; shaft and point join in a continuous arc; 
length 24 (12-27), width of base 12 (11-14). Dorsal anchor of same 
general structure as ventral, but with addition of well-defined roots; 
length 18 (16-20), vddth of base 9 (8-10). Bases of each anchor pair 
joined together by a transverse bar, the bars articulated to each other 
in an unusual fashion (see Discussion section for details). Ventral bar of 
simple construction (Fig. 5) and mildly curved throughout its length; 
length 37 (35-40). Dorsal bar consists of three fused portions: a simple 
midregion and with a shelf -Hke structure at each end (Fig. 4); length 
40 (37-42). Two bands of muscle tissue arise in the body proper, pass 
visibly through the peduncle and into the haptor, where each band at- 
taches to one of the shelf -like projections of dorsal bar ( Fig. 1 ) . Hooks 
14 in number (seven pairs), five pairs on ventral aspect of haptor, re- 
maining two pairs on dorsal aspect. Hooks similar in shape and size; 
each composed of ( 1 ) a solid, elliptical base, ( 2 ) a poorly differentiated 



ISIew trematode genus 



173 




Figures 1-8. Camera lucida illustrations of Gussevstrema amacleiih- 
rium n. gen., n. sp. 1, Whole mount, ventral view. 2, Ventral anchor. 3, 
Dorsal anchor. 4, Dorsal bar. 5, Ventral bar. 6, Hook. 7, Cirrus. 8, Ac- 
cessory piece. 

shaft, and (3) a sickle-shaped termination provided with an opposable 
piece (Fig. 6). Range of hook lengths 13 (12-14). 

Copulatory complex composed of a tubular cirrus arranged in a coil 
of ca. 1/4 turns, and a simple, basally articulated accessory piece (Figs. 
7, 8). Estimated length of cirrus 45; length of accessory piece 13 (12- 
14). Testis postovarian, much larger than ovary. Most of vas deferens ob- 



SMI!!1S0N!AN 

iNsinuTior; 



MAY 2 9 1969 



174 Proceedings of the Biological Society of Washington 

scured by the heavy vitellaria. Vagina and seminal receptacle not ob- 
served with certainty, however, a non-sclerotized vaginal opening is 
apparently present to right of median line, posterior to copulatory com- 
plex. 

Vitellaria very dense, the granules small and of uniform size. The 
granules become confluent: (1) in pharyngeal region, (2) between copu- 
latory complex and ovary, and (3) in posterior region of body proper, 
just anterior to peduncle. Intestinal crura difficult to observe because of 
vitellaria, but the strong posterior confluency of vitellarial granules 
suggests the hkelihood of a posterior confluency of the intestinal Hmbs. 

Discussion: The haptoral bars of this parasite are joined to each other 
in a unique way. The articulations are five in number. On the dorsal 
aspects of the bars, four articulations are seen: two occur near the mid- 
regions of the bars while an additional junction occurs distaUy, near 
either end. The fifth juncture, located on the ventral surface of the bars, 
is apparently composed of either muscle or connective tissue; this con- 
nective is irregular in shape. The bars are near the same level in the 
haptor, but are separated sufficiently so that the bars can be referred 
to as being either ventral or dorsal in the haptor. 

The haptor seems to be differentiated into two regions: a denser region 
located centrally, to which the anchors and bars attach, and a less dense 
region which we might term the "haptor proper." 

As pointed out by Price ( 1967 ) , only a few genera of the subfamily 
Ancyrocephalinae possess bars which are joined to each other. Gussev- 
strema is apparently most closely related to Actinocleidus Mueller, 1937 
and to Anchoradiscus Mizelle, 1941. The present genus, however, is 
differentiated from these two genera by the manner of articulation of 
the bars and by the position of the bars in reference to the haptor. Al- 
though Gussevstrema possesses only two eyespots, whereas members of 
both Actinocleidus and Anchoradiscus possess four, this reduced number 
of eyespots is not deemed worthy of generic status in the present case. 
This decision is based upon the presence of what are apparently com- 
prising granules in the anterior cephaHc region; these granules might be 
the dissociated elements of an additional vestigial pair of eyespots. 
Although we do not employ host-parasite relationships as criteria for 
establishment of new taxa, we believe that one aspect should be pointed 
out. Both Actinocleidus and Anchoradiscus are found on freshwater 
fishes native to North America. Terapon jarbua, on the other hand, is 
quite cosmopolitan in distribution and survives as readily in a marine 
environment as in fresh water. 

Etymology: The genus name is chosen to honor A. V. Gussev of the 
Zoological Institute of the Academy of Sciences of the U.S.S.R., in 
recognition of his work on monogenetic trematodes. The species name, 
amacleithrium, is a combination of two Greek words: amd, meaning 
"together" and cleithr-, meaning "bar," thus indicating the articulated 
nature of the haptoral bars. 



ISJew trematode genus 175 

Acknowledgment: Sincere thanks are extended to Miss Anne Dnicken- 
miller, of the Art Department of Millersville State College, for preparing 
the illustrations used in this study, and to Mr. T. Pike, Fisheries Re- 
search Officer of the Natal Parks, Game and Fish Preservation Board, 
Pietermaritzburg, Republic of South Africa, who donated the host speci- 
mens utilized in this study. 

Literature Cited 

Hargis, W. J., Jr. 1958. A revised, annotated Hst of terms useful for 
morphological studies of monogenetic trematodes. (Mimeo- 
graphed at Virginia Institute of Marine Science, Gloucester 
Point, Virginia). 12p. 

Price, C. E. 1966. Urocleidus cavanaughi, a new monogenetic trema- 
tode from the gills of the keyhole cichlid, Aequidens maroni 
(Steindachner). Bull. Georgia Acad. Sci., 24: 117-120. 

. 1967. Proposal of Syncleithrium, a new genus of the 

North American Monogenea. Texas J. Sci., 19: 175-183. 

, and H. P. Arai. 1967. A proposed system of anatomy for 

freshwater Monogenea. Canadian J. ZooL, 45: 1283-1285. 

, and T. E. McMahon. 1967. The monogenetic trematodes 

of North American freshwater fishes. Riv. Parassit., 28: 177- 
220. 



176 Proceedings of the Biological Society of Washington 



I 



? 

QH 

i 
B4X 

f^^ 82, pp. 177-188 29 May, 1969 

PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



STUDIES ON SPIROBOLOID MILLIPEDS. VII. 

A REMARKABLE NEW GENUS AND SUBFAMILY 

OF THE SPIROBOLELLIDAE 

FROM VERA CRUZ, MEXICO 

By Richard L. Hoffman 
Radford College, Radford, Virginia 

The milliped fauna of Mexico is, with continued exploration 
and study, rapidly assuming a position of world preeminence 
as regards its size and significance. The commingling of north- 
ern, tropical, and endemic elements has produced a diversity of 
novel and interesting millipeds that seems likely to yield ex- 
ceptional finds for many decades to come, and the present 
total of about 400 species known from the country will quite 
likely be increased tenfold before the end is reached. 

A major advance in our knowledge of Mexican diplopods 
was realized as one result of a prolonged collecting trip (July 
1965 to September 1966) by Dr. G. E. Ball and Mr. Donald R. 
Whitehead, of the Department of Entomology, University of 
Alberta. Although primarily concerned with the capture of 
carabid beetles, these two entomologists devoted their time 
and energies to the accumulation of various other arthropods, 
and have very generously presented me with a magnificent 
collection of several thousand well-preserved millipeds taken 
in many of the Mexican states. Preliminary sorting reveals that 
the majority of the species represented, and many of the genera, 
represent undescribed taxa, and obviously many years will 
elapse before the material can be adequately studied and the 
results integrated with our present knowledge of Central 
American millipeds. 

One specimen noted during sorting is of such special interest, 
however, that its immediate consideration is warranted as it 

12r— Proc. Biol. Soc. Wash., Vol. 82, 1969 (177) 



^RARIES 



178 Proceedings of the Biological Society of Washington 

seems to belong to the spiroboloid family Spirobolellidae, pre- 
viously unknown in Central America north of the Panama 
Canal Zone. The discovery of a new generic type in southern 
Vera Cruz is in itself important zoogeographically; moreover 
the species involved is of considerable interest because of sev- 
eral outstanding structural peculiarities. 

Family Spirobolellidae Brolemann 
Typhlobolellus new genus 

Type species: T. whiteheadi, new species. 

Diagnosis: A spirobolellid genus characterized by the absence of ocelli, 
by the origin of the ozopores on the 3rd segment, by the cristate modi- 
fication of the basal podomeres, by the formation of a sympleurotergal 
bridge behind the metasterna, and by the retention of sclerotized sternal 
structures in association with the phallopods. The first four characters 
mentioned are unique within the order SpiroboUda; the second, further- 
more, is unique within the Diplopoda. 

Distribution: Southern Vera Cruz, near sea level. 

Typhlobolellus whiteheadi new species 
Figures 1-8 

Holotype: Adult male, deposited in the USNM, from sea-level swamp 
forest, 0.8 miles west of Sontecomapan (ca. 11 miles north of Catemaco), 
Vera Cruz; 18 September 1965, G. E. Ball and D. R. Whitehead, col- 
lectors. 

Length about 28 mm. (specimen broken), body sHghtly compressed, 
transverse diameter ca. 1.3 mm., vertical diameter ca. 1.4 mm. at mid- 
body; general appearance remarkably slender for a spiroboloid and more 
similar to that of a cambaloid, the W/L ratio 4.9 percent. Body with 
55 segments, the last two legless. 

Head of the form shown in Figure 1, smooth and polished, with a few 
randomly dispersed facial setae and 2r-2 clypeal setae. No trace of ocelH. 
Parietal sclerite small, subtrangular, but clearly distinct. Basal segment 
of mandible moderate in size, vertically elongated; distal segment much 
smaller than usual for the order, extending less than half the length of 



Figs. 1-8. Typhlobolellus whiteheadi n. sp. 1, head and first six 
body segments, lateral view. 2, gnathochilarium. 3, ventral region of 
midbody segment, ventral aspect, legs removed; PS: presternum; MS: 
metasternum; PL: right pleuron; PTB: pleurotergal bridge isolating 
metastemum from caudal edge of segment. 4, posterior end of body, 
lateral aspect (legs not shown). 5, leg from midbody segment showing 
modification of prefemur and femur, and reduced setation. 6, coleopods, 



New spirohoioid millipede genus 



179 




posterior aspect. 7, coleopods, anterior aspect. 8, left phallopod and 
associated sternal structures, posterior aspect. Drawings from male 
holotype. Figures 1 and 3 originally made with magnification of 45 X, 
the others 90 X. 



SMintSONIAN Mflvoqiqcp 



180 Proceedings of the Biological Society of Washington 

genal edge. Gnathochilarium as figured (Fig. 2), no separate cardine 
sclerites observed. 

Antennae remarkably long and slender, extending back to 6th seg- 
ment when appressed to body, individual antennomeres cylindrical or 
very slightly compressed (especially distally), moderately clavate, the 
6th article largest and subfusiform; 7th small, discoid, with four minute 
terminal sensory cones. No special sensory setae or areas observed on 
other articles. 

CoUum (Fig. 1) smooth and pohshed, laterally broadly rounded and 
narrowly margined along anterior edge up to level of craniomandibular 
articulation; surface of lateral ends with several indistinct longitudinal 
striations. 

Segments of body essentially similar in structure; each prominently 
constricted at midlength producing two subsegments of virtually equal 
diameter, but since telescoping of the successive segments is only mod- 
erate, the body profile is distinctly submoniHform. Deepest part of each 
constriction with a sharp-edged transverse suture Hne marking anatom- 
ically the union of prosomite and metasomite, the latter apparently not 
subdivided into mesozonite and metazonite as in most other spiroboloid 
famihes. Traces of dorsolateral longitudinal sutures discemable just 
behind ozopores with suitable illumination; other segmental sutures ob- 
hterated. Surface of segments smooth and pohshed, microscopically reticu- 
lated; lower parts of metasomites with several indistinct longitudinal 
striations. Ozopores small but distinct, located at about the midlength 
of each metasomite of segments 3 through 54. 

Pleurosternal structure (Fig. 3) unusual in that the metasterna do not 
form the ventral part of caudal edge of segment, but are enclosed by 
medially projecting and coalesced elements of both the pleural sclerites 
and ventral ends of the metasomites, forming a "sympleurotergal" bridge 
similar to that of the 7th segment of males in other spiroboloid families. 
Anterior sternum slightly convex, finely and densely punctate; posterior 
sternum flat, smooth except for a few microscopic striations. Pleural 
sclerites smooth, their sutures clearly visible in dry material. 

Posterior segments ( Fig. 4 ) not especially telescoped, the last produced 
into a moderate-sized, cuneate epiproct which does not entirely conceal 
the paraprocts in dorsal aspect; latter strongly convex, smooth, the medial 
margins inflated and set off by prominent basal grooves, each with a 
marginal series of 11 or 12 conspicuous setiferous tubercles. Hypoproct 
transversely triangular, flat and smooth, without peculiarities. 

Legs (Fig. 5) short, not visible from above when extended laterally, 
differing from those of all other known spiroboloids in that the preferpora 
and femora are strongly convex dorsally, with a median row of about 
six acute dechvent spines. Ventral setae of midbody legs reduced: 1-1-1- 
1-1-1, usually a small dorsal seta at end of tarsal segment and an even 
smaller one ventrally at base of tarsal claw. Anterior legs without coxal 
lobes or other modifications, the tarsal claws elongate and slender. 

Ventral side of 7th segment continuous across a sympleural bridge, but 



New spiroboloid millipede genus 181 

without any modification of the median, postgonopodal area, the surface 
merely flat and undistinguished. Gonopods basically similar to those of 
Spirobolellus and related genera. Coleopods (Figs. 6, 7) with well- 
defined transverse sternum, medially produced into an elongate, digitiform 
process; coxae with prominent deep notch along ventrolateral edge in 
front of lateral articulation of telopodite; telopodites of moderate size, 
separated on the posterior side from the median prolongation of the 
sternum by a narrow lobe of the coxa; distally lobed, each lobe with a 
subterminal row of fine setae along the outer edge. 

Phallopods (Fig. 8) similar to those of other spirobolelHds in general 
form, attached to a delicate but prominent sternal structure, heretofore 
not recorded for this family; gonopod divided by an oblique suture into 
a broad, subquadrate coxal remnant and a more slender and elongated 
telopodite, the latter also divided (secondarily?) into a basal slender 
trunk and a broad, laminate, triangular distal lobe set at right angles. 
Coxa with slender apodemes attached at midlength of its ventral edge as 
usual for the family. No trace of grooving or glandular modification 
noted in the thin, flattened, and delicate phallopods. 

Color pattern faded through preservation, but generally pale testa- 
ceous brown with metasomites distinctly darker, imparting an annulated 
appearance, the pigmentation more intense on caudal third of body; 
last segment and paraprocts nearly blackish. Head, antennae, legs, and 
lower sides pale yellow. 

General Observations 

1. Trypsin as a clearing agent. The small size of the type specimen 
of T. whiteheadi and obvious dehcacy of its structure required special 
techniques for preparation prior to study. The use of KOH as a macer- 
ating material, while suitable for large and robust diplopods, seemed too 
hazardous in this case, so that removal of muscle tissue and internal 
organs was accomplished by breaking the specimen between body seg- 
ments 7 and 8, and soaking the anterior end of the body overnight in an 
aqueous solution of commercial C.P. trypsin at a temperature of approxi- 
mately 100° F. The use of this technique is recommended to the atten- 
tion of other students of small and fragile arthropods. All of the non- 
sclerotized tissue can be digested and washed out without any damage 
to even the most delicate sclerotized structures, something that can not 
be asserted for caustic agents even with dilute solutions and moderate 
warming. 

2. Phallopod sternum. The discovery of a fairly extensive sternal rem- 
nant to which the posterior gonopods are attached is of considerable 
interest from the standpoint of phylogeny. Unfortunately the structure 
was unknowingly damaged during dissection, so that the drawings show 
a somewhat distorted appearance. There seems to be both transverse 
elements and proximally directed apodemes (= tracheal apodemes?), 
and if the latter possibility can be confirmed by future studies, a new 
light will be shed upon the homology of the phallopod basal apodemes 



182 Proceedings of the Biological Society of Washington 

in other spiroboloid groups. The presence of sternal elements connected 
with the phallopods, of course, can only be considered as plesiomorphic 
in character. 

3. Notes on the classification of the Spirobolellidae. When first set up 
by Brolemann in 1913, this family contained only three genera. In my 
opinion, there are now no less than 16 nominal generic taxa here refer- 
able, many of course being of quite dubious validity. 

Although Brolemann defined his family groups with considerable pre- 
cision, later workers either ignored the family category in the SpiroboUda 
altogether (Chamberlin, Verhoeff), or combined the SpirobolelHdae 
with the Spirobolidae (Attems, Carl). As there has been no synopsis of 
the family since 1914, it certainly seems important to list here the 
generic names that have been based upon taxa agreeing with Brole- 
mann's diagnosis and to propose some disposition of their taxonomic 
status. Full hterature citations may be extracted from the recent hst 
(Hoffman & Keeton, 1960) of generic names in the Spirobolida. 

Aporobolus Loomis, 1934 Tobago 

Attemsobolus Verhoeff, 1924 Australia 

Barrobolus ChamberHn, 1925 Panama 

Carlobolellus Brolemann, 1931 New Caldonia 

Desmocricellus Attems, 1953 New Zealand 

Howeobolus Verhoeff, 1928 Lord Howe Island 

Mauritobolus Verhoeff, 1939 Mauritius (native?) 

Microspirobolus Silvestri, 1898 Venezuela 

Paraspirobolus Brolemann, 1902 Brasil 

Physobolus Attems, 1936 India 

Poratobolus Verhoeff, 1924 Austraha 

Queenslandobolus Verhoeff, 1924 Australia 

Sechellobolus Brolemann, 1913 Seychelles (native?) 

Spirobolellus Pocock, 1894 Sumatra 

Spirobolinus Silvestri, 1898 Ecuador 

Walesbolus Verhoeff, 1928 Austraha 

The taxonomy of the foregoing ensemble is at present quite unsatis- 
factory. Brolemann's concept of Spirobolellus was based upon S. rainbowi 
Brol., an AustraHan species, whereas the type of the genus, S. chrysodirus 
Pocock, was from Celebes and its gonopod structure was not accounted 
or figured. Inferentially, however, it seems highly probable that the 
two species are congeneric, as there is now known a fairly large group 
of similar species ranging from Sumatra to Micronesia and south to New 
Caledonia, Austraha, and New Zealand. J. Carl ( 1926 ) has given a good 
account of the Caledonian fauna and remarked, quite correctly in my 
opinion, that many if not all of Verhoeff 's 1924 generic names are really 
based only on species characteristics. What Carl did not take into ac- 
count, nor has anyone else more recently, is that a generic separation 
between the Indoaustrahan taxa referred to Spirobolellus, and those of 
the New World to Microspirobolus, appears impossible. Aside from 



New spiroboloid millipede genus 183 

literature accounts, I have compared some specimens from the Carohne 
Islands with Haitian material, and can find no justification for the recog- 
nition of two genera. 

The name Microspirobolus has been used for a large number of Neo- 
tropical, chiefly Antillean, species, but here again the type species was 
described without reference to its gonopod structure so that the exact 
status of M. pulchellus remains a little dubious. But in this case, Silvestri 
later asserted that the two new species M. marmoratus and M. insularis 
(1908), which were well-described and illustrated, are congeneric with 
pulchellus, and Silvestri rarely if ever overlooked possible generic dis- 
tinction. 

The generic name Barrobolus has already been combined with Micro- 
spirobolus by Loomis ( 1964 ) . I have briefly considered the status of 
the two names Aporobolus and Spirobolinus in 1955 without reaching 
a definite conclusion; both nominal taxa lack ozopores on the 7th seg- 
ment but are otherwise typical spirobolellids. I here retain the two names 
but have no illusions about a long life for either of them. 

Paraspirobolus was based upon immature males of a species that has 
not been reported since 1902 and remains something of an enigma. 
Schubart ( 1947 ) suggests that P. paulistus Brol., the type and only spe- 
cies, was based upon specimens of Sechellobolus dictyonotus (Latzel) 
which is synanthropically not rare in southern Brasil. I agree entirely 
with this possibility as there is no reason to believe that an endemic 
spirobolellid genus occurs in that part of South America. Sechellobolus 
(misspelled Seychellobolus by several authors) itself seems to contain 
only its type species S. dictyonotus (Latzel), which is known from Berlin 
and Hamburg (the type locaHtyl), the Seychelles, Mauritius, and 
Brasil. I have personally identified specimens also from Jamaica, and 
beheve that the milHped described from Annam by Graf Attems ( 1953 ) 
as Physobolus striatus is the same form. In view of such a far-flung dis- 
tribution, the relative paucity of records is surprising, and the original 
home of the species has yet to be ascertained. 

Assuming the probable correctness of the synonymies mentioned 
above, we find that Paraspirobolus is the older of the two generic syno- 
njntns, while dictyonotus (Latzel, 1895) is the oldest of several specific 
names that seem to have been based upon the same species. In most 
respects this genus is similar to Spirobolellus but can be distinguished 
on the basis of what appears to be a receptacle (seminal?) in the telop- 
odite of the phallopod. 

Attemsobolus is likewise weakly differentiated, its type species is 
marked chiefly by the presence of a secondary process on the coleopod 
telopodite (Verhoeff wrote that the telopodite was deeply bifidl). 
Howeobolus and Walesbolus are not well-described, the gonopods ap- 
parently having been torn apart before the drawings were made, but 
the names may be retained provisionally. 

Physobolus, however, appears to be a well characterized genus with 
two species in the "Farther India" region. The sternum of the colepods 



184 Proceedings of the Biological Society of Washington 

is small and transverse, leaving the coxal plates entirely exposed; the 
phallopod seems to be two-segmented with a distinct gland located in 
the basal (coxal?) division. 

Taking the foregoing remarks collectively, it is possible to render the 
list of generic names into a somewhat more condensed roster, and to 
align them in three groups which can be considered as subfamilies for 
the present. I believe that the suggested subordination of six names 
constitutes new synonymy in each case. Characteristics of the subfamilies 
can be readily derived from those of the included genera, the Spirobolel- 
linae for instance includes all of the species that lack the specialized 
features of Physobolus and Typhloholellus. Of these last two genera, 
Typhlobolellus is by far the more disjunct, and may well warrant family 
status in the light of future investigations. 

The proposed classification may be rendered thus: 

FAMILY SPIROBOLELLIDAE BROLEMANN, 1913 
Subfamily Spirobolellinae, n. subfam. 

Spirobolellus Pocock, 1894 (syns. Barrobolus Chamberlin, 1925, 
Mauritobolus Verhoeff, 1939, Microspirobolus Silvestri, 1898, Poro- 
tobolus Verhoeff, 1924, Queenslandobolus Verhoeff, 1924). East 
Indies, Micronesia, New Caldonia, eastern Australia; Panama, 
Colombia, Ecuador, Peru, Venezuela, West Indies (except Ja- 
maica! ) . 

Aporobolus Loomis, 1934. Tobago. (= Spirobolinus?). 

Attemsobolus Verhoeff, 1924. Queensland. 

Carlobolellus Brolemann, 1931. New Caledonia. 

Desmocricellus Attems, 1953. New Zealand. 

Howeobolus Verhoeff, 1928. Lord Howe Island. 

Paraspirobolus Brolemann, 1902 (syn. Sechellobolus Brolemann, 
1913). Widely dispersed by commerce, perhaps endemic in south- 
east Asia. 

Spirobolinus Silvestri, 1898. Ecuador. 

Wdesbolus Verhoeff, 1928. New South Wales. 
Subfamily Physobolinae, n. subfam. 

Physobolus Attems, 1936. Northern India; Vietnam. 
Subfamily Typhlobolellinae, n. subfam. 

Typhlobolellus, n. gen. Southern Mexico. 

4. Some notes on distribution and relationships. The presently known 
distribution of the Spirobolellidae is especially interesting; in the main 



Fig. 9. Distribution of the family SpirobolelHdae (broken line) 
and the family Atopethohdae (cross-hatching). The type locality of 
Typhlobolellus whiteheads is indicated by the black spot in southern 
Mexico. Records for the family in the Indian Ocean ( Seychelles, Mauri- 
tius) are presumed to be due to accidental introduction through com- 



New spiroboloid millipede genus 



185 



» 




merce. The two areas circled on the mainland of southeast Asia indicate 
the occurrence of the two known species of Physobolus and the sub- 
family Physobolinae. With the foregoing exceptions the distribution of 
Spirobolellus is virtually co-extensive with that of the family. 



186 Proceedings of the Biological Society of Washington 

points it agrees closely with that of the Rhinocricidae although less ex- 
tensive. The map (Figure 9) shows in a general way the occurrence of 
spirobolellids as here understood. The greatest numbers of species are 
to be found on islands, such as New Caledonia (25) and Hispaniola 
(10), remarkably few are so far known from continental regions (south- 
east Asia, northern South America). Perhaps this reflects a dechning 
evolutionary status for the group. Geographic ranges shared between 
the East and West Indies are not to be explained by recourse to "drift" 
or other postulated crustal movements unless we can suppose that an- 
cestral forms occurred widely across a single northern continent ("Laur- 
asia") in the Palaeozoic, prior to the formation of the Atlantic Ocean 
and prior to contact between the two subsequent northern landmasses 
with South America and Africa, respectively. 

As already pointed out some years ago (Hoffman & Orcutt, 1960, p. 
112), the family most closely related to the Spirobolelhdae appears to 
be the North American endemic group AtopethoHdae, the present range 
of which is shown by cross-hatching on the map. On the basis of gonopod 
structure, one might suspect that the atopethoHds are perhaps somewhat 
more primitive, but speculations of this sort are still quite premature. 

Doubtless the Rhinocricidae is to be regarded as belonging to the 
same group of families. Although rhinocricids do extend into Brasil and 
Argentina, all of the far-southern species are quite homogeneous and 
the center of diversification in this family is located in northern South 
America and the West Indies. Antipodal vicariation at the generic level 
is to be noted between Rhinocricus (Haiti, Porto Rico, Cuba) and Acla- 
docricus ( Indoaustrahan region generally), and between Neocricus (Ven- 
ezuela, Colombia) and Dinematocricus (Australasia). 

One inference to be drawn from the foregoing remarks is that the two 
families Rhinocricidae and Spirobolelhdae originated in the northern 
hemisphere and have invaded South America (during the late Mesozoic 
or, perhaps, later, early Pliocene) only very shghtly; neither group is 
represented in temperate eastern Asia, the Indian peninsula, or in Africa. 
Probably the present distribution of spirobolellids is due to their virtual 
displacement on continental areas by members of more modern and/or 
successful famihes (e.g., AtopethoHdae and Rhinocricidae), although 
the mechanics of competition and displacement among milliped groups 
have yet to be demonstrated factually. 

Literature Cited 

Attems, C. 1953. Myriopoden von Indochina. Expedition von Dr. C. 

Dawydoff (1938-1939). Mem. Mus. Nat. hist, natur., ser. A, 

vol. 5, pp. 133-230, figs. 1-119. 
Brolemann, H. W. 1902. Myriapodes du Musee de Sao Paulo, Rev. 

Mus. PauHsta, vol. 5, pp. 35-237, figs. 1-271. 
. 1913. Un nouveau systeme de spirobolides. Bull. Soc. Ent. 

France, no. 19, pp. 476-478. 



"New spiroboloid millipede genus 187 

. 1914. Etude sur les spirobolides. Ann. Soc. Ent. France, 

vol. 83, pp. 1-38, figs. 1-9. 

Carl, J. 1926. Diplopoden von Neu-Caledonien und den Loyalty- 
Inseki, in: Sarasin & Roux, ed., Nova Caledonia, Zool., vol. 
IV, pp. 369-462, figs. 1-159. 

Hoffman, R. L. 1955. Studies on spiroboloid millipeds. III. The 
genus Spirobolinus Silvestri. Proc. Biol. Soc. Washington, 
vol. 68, pp. 151-154, figs. 1-4. 

AND B. S. Orcutt. 1960. A synopsis of the Atopetholidae, 

a family of spiroboloid millipeds. Proc. U. S. Nat. Mus., vol. 
Ill, pp. 95-165, figs. 1-11. 

AND W. T. Keeton. 1960. A hst of the generic names pro- 
posed in the diplopod order Spirobolida, with their type spe- 
cies. Trans. American Ent. Soc, vol. 86, pp. 1-26. 

LooMis, H. F. 1964. The milHpeds of Panama (Diplopoda). Fieldi- 
ana: Zool, vol. 47, pp. 1-136, figs. 1-13. 

Silvestri, F. 1908. Myriopoda from Porto Rico and Culebra. Bull. 
American Mus. Nat. Hist., vol. 24, pp. 563-578, figs. 1-11. 

ScHUBART, O. 1947. O elemento "synanthropo" e estrangeiro entre os 
Diplopoda do Brasil. Arthropoda, vol. 1, pp. 23-40. 



188 Proceedings of the Biological Society of Washington 



L 82, pp. 189-192 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 

THE TAXONOMIC STATUS OF MANDIBULOPHOXUS 
GILESI BARNARD, 1957 (CRUSTACEA: AMPHIPODA) 

By W. Scott Gra.y, Jr. and John C. McCain 

University of the Pacific, Pacific Marine Station, Dillon Beach, 

California; Smithsonian Oceanographic Sorting Center, 

Washington, D. C. 

Barnard (1957) erected the genus Mandibulophoxus to ac- 
commodate those species of phoxocephalids in which the man- 
dibular palp was attached to a bulky process on the body of 
the mandible. At this time he described the new species, Man- 
dibulophoxus gilesi, from a single specimen and named it as 
the type-species of the genus. In his excellent monograph on 
the eastern Pacific Phoxocephalidae (1960) he synonymized 
M. gilesi with M. uncirostratus (Giles, 1890) on the basis of 
Pillai's ( 1957 ) paper. Specimens that agree well with Barnard's 
description of M. gilesi have been collected in Tomales Bay, 
California and Yaquina Bay, Oregon and they are herein com- 
pared with the descriptions of M. uncirostratus by Giles and 
PiUai. 

Mandibulophoxus gilesi Barnard, 1957, new synonymy 

Mandibulophoxus gilesi Barnard, 1957, pp. 433-435, figs. 1-2. 

Mandibulophoxus uncirostratus (not Giles, 1890). — Barnard, 1960, p. 
359 [in part]. 

Material examined: California, Tomales Bay, offshore from Pacific Ma- 
rine Station, 2.1 meters in fine sand, 2 $ $ (3.5-4.7 mm). 
Oregon, Yaquina Bay, near oceanic end of south jetty. 5.5-7.3 meters 

in coarse sand, 5 $ $ (3.7-5.2 mm). 

Discussion: A comparison of the eastern North Pacific specimens with 
the description and figures of Pillai (1957) has revealed the following 
differences: 

1. Pillai's figures of M. uncirostratus show fewer spines on the rami of 
uropods 1 and 2 than on the eastern North Pacific specimens. His figure 
V-12 of uropod 1 show 4 spines on both rami whereas our specimens 

13— Prcx:. Biol. Soc. Wash., Vol. 82, 1969 ( ] 



f JUN2bi969 j 



190 Proceedings of the Biological Society of Washington 




Figure 1. Mandibulophoxus gilesi from Yaquina Bay. a, telson 
b, lateral view; c, gnathopod 2; d, gnathopod 1; e, uropod 1; /, mandible 
g, pereopod 1; h, uropod 2; i, m-opod 3; /', pereopod 2; k, pereopod 3 
I, pereopod 5; m, maxilla 1; n, maxilla 2. 



have 5 on the inner ramus and 6 on the outer. Pillai's fig. V-13 shows 
3 spines on the inner ramus of uropod 2 and 4 on the outer. Our speci- 
mens have 5 spines on the inner ramus of uropod 2 and 7 on the outer. 
2. Pillai's figure V-11 shows 3 or 4 teeth on article 2 of pereopod 5 
while our eastern North Pacific specimens have over 6 teeth. In addi- 
tion article 4 is figured as much longer than article 5 but they are al- 
most equal in our specimens. 



Taxonomic status of Mandibulophoxus 191 

3. Giles (1890) and Pillai both state and figure the rostrum of M. 
uncirostratus as deflexed or hook-shaped; in our specimens the rostrum 
is only shghtly bent downward. 

4. Pillai states that M . uncirostratus has a tooth in addition to a strong 
spine on the palm of gnathopods 1 and 2. In fig. V-6 he shows a well 
defined sharp cusp on the distal posterior margin of the palm of the 
propodus, presumably the tooth to which he was referring. Our speci- 
mens bear no tooth on the palm of the gnathopods but do have what 
might be considered a strong spine and a well defined but rounded cusp. 
The proportions of the articles of gnathopod 2 are unusual in Pillai's 
figures but this is undoubtedly due to the omission of the joint between 
articles 4 and 5. 

5. Pillai states and his figures show that the inner ramus of uropod 
3 is small, less than % the length of the first segment of the outer ramus. 
In our specimens the inner ramus is almost equal in length to the first 
segment of the outer ramus. 

6. Pillai's figures of pereopod 3 of M. uncirostratus show that article 4 
is shorter than article 5; in our specimens these articles are approximately 
equal in length. 

7. The process on the body of the mandible to which the mandibular 
palp is attached was figured by Pillai as being much shorter and less 
distinct in M. uncirostratus than in our specimens. 

8. In Pillai's figures of M. uncirostratus article 5 of pereopod 4 is ap- 
proximately V2 to % the lengths of articles 4, 6, and 7; in our specimens 
articles 4, 5, and 6 are approximately equal in length and article 7 is ap- 
proximately % the lengths of articles 4, 5, and 6. 

Other differences between Pillai's figures and our specimens, particu- 
larly in the proportions and degree of setation of various appendages 
could be merely artifacts of illustration and we do not beheve their value 
to be as great as the above-mentioned differences. Pillai's figures show a 
convex dorsal outhne in lateral view, whereas all our specimens were 
concave. Whether this curvature varies with the type of preservation or 
is a true difference between the species requires examination of living 
specimens. 

In view of the differences Hsted above, we are reinstating Barnard's 
species, M. gilesi. M. gilesi thereby has a known distribution from Oregon 
to Southern Cahfomia and M . uncirostratus is hmited to the Madras and 
Ceylon coasts. 

Species of Paraphoxus show sexual dimorphism in the inner ramus of 
uropod 3 which is relatively longer in the male. As far as we know, males 
of M . gilesi and M. uncirostratus have not been described. Giles does not 
state the sex of his specimen. 

Literature Cited 

Barnard, J. L. 1957. A new genus of phoxocephahd Amphipoda 
(Crustacea) from Africa, India, and California, Ann. Mag. 
Nat. Hist., ser. 12, vol. 10, pp. 432-438, 4 figs. 



SMI!^!S0NIA^ 

iK'siiTuimr 



MAY 2 9 1969 



192 Proceedings of the Biological Society of Washington 

. 1960. The amphipod family Phoxocephalidae in the east- 
em Pacific Ocean, with analyses of other species and notes 
for a revision of the family. Allan Hancock Pacific Exped., 
vol. 18, no. 3, pp. 175-368, 75 pis. 

Giles, G. M. 1890. Descriptions of seven additional new Indian 
amphipods. Journ. Asiatic Soc. Bengal, vol. 59, pt. 2, pp. 
63-74, pi. 2. 

PiLLAi, N. K. 1957. Pelagic Crustacea of Travancore. III. Amphi- 
poda. Travancore Univ., Bull. Central Res. Inst., ser. C, vol. 
5, no. 1, pp. 29-68, 18 figs. 



82, pp. 193-200 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



A NEW CRAWFISH OF THE GENUS HOBBSEUS 
FROM MISSISSIPPI (DECAPODA, ASTACIDAE) 

By Joe B. Black 

Department of Biological Sciences, 

McNeese State College, 

Lake Charles, Louisiana 

The species described herein is assigned to the genus Hobb- 
sens of the crawfish subfamily Cambarinae. The members of 
this newly designated genus (Fitzpatrick and Payne, 1968: 15) 
formerly constituted the Cristatus Section of the genus Cam- 
harus (Hobbs, 1955: 95). Other described species in the genus 
include H. cristatus (Hobbs, ibid.) from the western Tombig- 
bee and Pascagoula River watersheds in Lowndes, Noxubee, 
Kemper and Lauderdale Counties, Mississippi; H. prominens 
(Hobbs, 1966: 110) from the eastern Tombigbee River water- 
shed in Sumter County, Alabama; H. valleculus (Fitzpatrick, 
1967: 163) from the upper Pearl River watershed in Choctaw 
County, Mississippi; and H. orconectoides Fitzpatrick and 
Payne (1968: 17) from the western Tombigbee River water- 
shed in Oktibeeha County, Mississippi. The present species 
has been collected from five localities in Winston County, 
and one locality in Neshoba County, Mississippi, all from the 
Pearl River watershed. 

Hobbseiis attenuatus new species 
Diagnosis: Body pigmented; eyes well developed and pigmented; ros- 
trum ovate and subspatulate, without marginal spines or tubercles. 
Areola 5.3-7.5 times longer than broad and constituting 30.0-34.2 per- 
cent of total length of carapace. Suborbital angle almost obsolete. Post- 
orbital ridges without spines. Antennal scale slightly less than one-half 
as broad as long. Chela vdth cristiform row of tubercles on mesial 
margin of palm. First pleopod of first form male with central projection 
directed at approximately 90 degree angle to main shaft of appendage; 

14 — ^Proc. Biol. Soc. Wash., Vol. 82, 1969 (193) 

JUN2D 1969 

sl'BRARlE^ 



194 Proceedings of the Biological Society of Washington 

mesial process extremely slender, directed at 90 degree angle, extending 
caudad approximately 40 percent of its length beyond tip of central 
projection. Pleopods symmetrical (Fig. 3). Sternum of first form male 
with conspicuous, ventrally projecting, setiferous processes at bases of 
third and fourth pereiopods. Annulus ventrahs freely movable. 

Holotypic male. Form I: Body subovate, compressed laterally. Ab- 
domen narrower than cephalothorax (7.5 and 8.5 mm in widest parts, 
respectively). Width of carapace less than depth at levels of greatest 
dimension (8.5-9.0 mm). Areola (Fig. 8) moderately narrow (6.1 times 
longer than wide), with row of punctations mesial to each branchio- 
cardiac groove and medial longitudinal shallow row in cephalic portion. 
Length of areola 31.3 percent of length of carapace. Rostrum relatively 
narrow (width 70 percent of length), subspatulate, with small up- 
turned acumen; margins sHghtly elevated; no lateral spines present; tip 
of rostrum reaching tip of distal end of peduncle of antennule; upper 
surface only slightly depressed, deepest in caudal portion of rostrum 
(Fig. 8); subrostral ridges weak but present. 

Postorbital ridges well developed; cephalic ends lacking tubercles or 
spines; suborbital angle, strongly obtuse, almost obsolete. Branchiostegal 
spine small but acute. Cervical spines and tubercles absent. Carapace 
punctate dorsally and cephalolaterally, punctations particularly con- 
spicuous in area immediately mesial to postorbital ridges; lateral por- 
tions of branchiostegites granulate. Abdomen longer than carapace (20.5 
and 19.5 mm). Cephahc section of telson with two acute spines in each 
caudolateral corner. 

Epistome (Fig. 11) subtriangular with elevated margins, lateral auri- 
cles, and small cephalomedian extension. Antennules of usual form with 
small spine on lower surface of basal segment. Antennae extend caudad 
to fourth abdominal segment. Antennal scale (Fig. 12) broadest distal 
to midlength, terminating in strong spine. 

Chela (Fig. 7) with broad palm, slightly inflated, length of inner 
margin of palm slightly greater than width ( 6.0-5.8 mm ) ; dorsal surface 
of palm with numerous setiferous squamous tubercles; lower surface of 
palm punctate; inner margin with cristiform row of 16 tubercles; fingers 
not gaping. Upper and lower surfaces of both fingers with submedian 
ridge flanked by setiferous punctations, ridges on upper surfaces more 
prominent; opposable margin of dactyl with 2 subequal tubercles along 
proximal one-half, distalmost largest, with crowded minute denticles 
along distal one-half; opposable margin of immovable finger with 2 sub- 
equal tubercles along proximal one-half and crowded minute denticles 
along distal one-half. 

Carpus of cheliped longer than broad; grooved dorsally, with scat- 
tered setiferous punctations, inner margin with numerous cristiform 
tubercles and one strong acute spine near distal end; lower surface with 
strong acute spine on distolateral margin. Merus with row of 13 small 
acute spines along entire lower mesial margin; lower lateral margin with 
irregularly spaced row of 10 small acute spines along proximal two- 



ISIev) Mississippi crawfish 



195 




Figures 1-12. Hobbseus attenuatus. (1) mesial view of first pleo- 
pod of holotype; (2) mesial view of first pleopod of morphotype; (3) 
caudal view of first pleopods of holotype; (4) lateral view of first pleo- 
pod of morphotype; (5) lateral view of first pleopod of holotype; (6) 
lateral view of carapace of holotype; (7) carpus and chela of holotype; 
(8) dorsal view of carapace of holotype; (9) ischiopodite and basipodite 
of third pereiopod of holotype; ( 10 ) annulus ventrahs and sternal plate 
of allotype; (11) epistome of holotype; (12) antennal scale of holotype 
(setae omitted from all figures). 



^iXSil? MAY 2 9 1969 



196 Proceedings of the Biological Society of Washington 

thirds, and four larger spines distally. Mesial margin of ischiopodite 
with row of six small acute spines. 

Ischiopodite of third pereiopod (Fig. 9) with strong simple hooks, 
extending proximally well beyond bases of ischiopodites. Coxae of fourth 
pereiopods without conspicuous prominence, those of fifth with ventro- 
mesial tubercle and more conspicuous one located cephalodorsal to the 
first and in contact with gonopod. Sternal projections at bases of cox- 
opodites heavily setiferous and conspicuous, forming deep trough for 
gonopods. 

First pleopod (Figs. 1, 3, 5) reaching to coxopodites of third pereio- 
pods when abdomen flexed, lying deeply between ventrally projecting 
sternal projections; distal portion terminating in two distinct parts; cen- 
tral projection corneous, tip rounded, bent caudally at angle of approx- 
imately 90 degrees to main shaft of appendage; mesial process arising 
from conspicuous knob near base, non-corneous, attenuate and parallel 
to central projection; mesial process conspicuously longer than central 
projection. 

Morphotypic male. Form II: Differs from holotype in following re- 
spects: tip of rostrum rounded, without tubercular acumen; areola pro- 
portionally narrower (6.5 times longer than wide); punctations in areola 
less conspicuous; tubercles along opposable margins of both fingers less 
prominent; usual differences occur in secondary sexual characters: re- 
duced and blunter stylets on gonopod, reduced hooks on third pereiopods 
and reduced prominences on coxae of fifth pereiopods. 

First pleopod ( Figs. 2, 4 ) not reaching base of third pereiopods, both 
processes non-corneous, less attenuate; directed at angles slightly less 
than in holotype; juvenile suture near base prominent; mesial process 
proportionally shorter. 

Allotypic female: Rostrum as in holotype with distinct tuberculate 
acumen. Differs from holotype in following respects: Areola proportion- 
ally wider (5.5 times longer than wide); epistome with less distinct 
auricles; cristiform tubercles on inner margin of palm less distinct and 
fewer in number ( 14 ) . 

Aimulus ventralis (Fig. 10) freely movable; median S-shaped sinus 
originating in cephahc depression sHghtly sinistral to median hne; sec- 
ondary depression originating from cephahc one-third of sinus extending 
caudosinistrally between two elevations. Sternal plate immediately caudal 
to annulus with median depression bordered by slight elevations. 

Measurements: as follows ( in mm ) : 

Holotype Allotype Morphotype 

Carapace 
Length 
Width 
Height 



19.5 


16.2 


16.0 


8.5 


7.5 


7.5 


9.0 


8.0 


8.0 



New Mississippi crawfish 197 







Holotype 


Allotype 


Morphotype 


Areola 










Length 




6.1 


5.5 


5.2 


Width 




1.0 


1.0 


0.8 


Rostrum 










Length 




5.0 


4.1 


4.0 


Width 




3.5 


3.1 


3.1 


Antennal scale 










Length 




4.2 


3.5 


3.5 


Width 




2.0 


1.7 


1.7 


Chela 










Length (lateral 


mar. ) 


12.5 


7.0 


8.9 


Palm length (mesial mar.) 


6.0 


3.8 


4.3 


Palm width 




5.8 


3.0 


3.8 


Dactyl length 




6.9 


4.2 


4.9 



Type locality: Roadside ditch adjacent to Noxapater Creek, on gravel 
road two miles north of State Route 395, six miles west of Noxapater, 
Winston County, Mississippi (R HE, T 14N). The ditch contains water 
in all but severe drought conditions and has abundant submergent and 
emergent aquatic vegetation. Dominant trees in the adjacent swampy 
area were Liquidamber, Pinus, Acer, Liriodendron. All specimens were 
captured with dipnets from open water. Other crawfishes associated 
with Hobbseus attenuatus in the ditch were Procambarus a. acutus 
(Girard, 1852:91), P. planirostris Penn (1953:71), Cambarus hedgpethi 
Hobbs (1948:224). At other locaHties the associates were P. a. acutus 
and C. d. diogenes Girard (1852:88). 

Disposition of types: The holotype, allotype, and morphotype are 
deposited in the United States National Museum ( Nos. 129535, 129536, 
and 129537, respectively). Paratypic series are in the United States Na- 
tion Museum (1^1, 2^ 5 II, 2? $ ), the Museum of Comparative Zo- 
ology (1^1, l^II, 19), Tulane University Invertebrate Collection 
(14^ ,^1, 4^ ^11, 262 $ ), and the Mississippi State University Collec- 
tion (151, 2$ 511, 2$ 9 ). Other paratypes (2$ $1, 135 511, 30$ 9, 
25 5 juv., 39 9 juv.) are retained in my personal collection. 

Ecological and life history notes: Presumably H. attenuatus, like other 
members of the genus, is a secondary burrower. All specimens came 
from temporary aquatic situations. Breeding form males have been col- 
lected in June (17 total), August (2) and December (1). Medium 
sized juveniles were collected in late December. 

Color notes: Primary background color of carapace ohve-brown with 
darker flecks of brown on dorsal surface becoming larger blotches on 
lateral surfaces. Base color of abdomen medium brown tinged with 
pink, pink tinge becoming more pronounced laterally and caudally. Mid- 
dorsal area darker brown, giving appearance of poorly defined stripe 



198 Proceedings of the Biological Society of Washington 

Each abdominal segment with rectangular patch of dark brown along 
cephalic edge of either side of mid-dorsal line, giving overall appear- 
ance of pair of dark brown stripes. Below rectangular patches is lightly 
marked pinkish-brown stripe of approximately 1.0 mm diameter, bor- 
dered laterally by weU defined stripe of dark brown. Dorsal surfaces 
of the chela pinkish-brown with oHve-brown blotches. 

Variations: Variations in number and size of tubercles on inner margin 
of palm were observed, the range being 13-17, males usually having a 
larger number than females. Several specimens were noticed to be with- 
out an acumen, in addition to the morphotypic male. The areola varies 
in width from 5.3-7.5 times longer than broad. 

Relationships: H. attenuatus is most closely related to H. cristatus 
(Hobbs, 1955:95). It is similar to the latter in the overall appearance 
and degree of flexion of the pleopods. Closer examination of the pleo- 
pods, however, reveals characteristic differences. The first pleopod of 
H. cristatus shows an extremely broad, flattened mesial process, less 
flexion of both mesial process and central projection, and a sharp tipped 
central projection. The annuli ventrales of the two species are quite 
dissimilar in sculpturing and curvature of the medial sinus. The epistome 
most closely resembles H. valleculus ( Fitzpatrick, 1967:163); the an- 
nulus ventralis is similar in many respects to H. orconectoides Fitzpatrick 
and Payne (1968:17). The most distinguishing feature of the new spe- 
cies, however, is the extremely long attenuate stylets of the first form 
male gonopod, together with the deep trough formed by the sternal 
projections to house the attenuate pleopods. 

Etymology: The name of this species is taken from the Latin attenuo, 
to make thin; it is so named because of the extremely thin mesial proc- 
ess of the first pleopod of the first form male. 

Acknowledgments: I am grateful to Drs. Horton H. Hobbs, Jr. of 
the Smithsonian Institution and J. F. Fitzpatrick, Jr. of Mississippi State 
University for loan of specimens of related species, to my wife and daugh- 
ter for assistance with field collections, and to Mr. Jerry G. Walls for 
critical review of the manuscript. 

Literature Cited 

Fitzpatrick, J. F., Jr. 1967. A new crawfish of the Cristatus Section 
of the genus Cambarus from Mississippi (Decapoda, Asta- 
cidae). Proc. Biol. Soc. Washington 80: 163-168. 

AND J. F. Payne. 1968. A new genus and species of craw- 
fish from the southeastern United States (Decapoda, Asta- 
cidae). Proc. Biol. Soc. Washington 81: 11-22. 

GiRARD, C. 1852. A revision of the North American Astaci, with ob- 
servations on their habits and geographical distribution. Proc. 
Acad. Nat. Sci. Philadelphia 6: 87-91. 

Hobbs, Horton H., Jr. 1948. A new crayfish of the genus Cambarus 
from Texas, with notes on the distribution of Cambarus 
fodiens (Cottle). Proc. U. S. Nat. Mus. 98: 223-231. 



'New Mississippi crawfish 199 

. 1955. A new crayfish of the genus Cambarus from Missis- 
sippi. Proc. Biol. Soc. Washington 68: 95-100. 

. 1966. A new crayfish from Alabama with observations on 

the Cristatus Section of the genus Cambarus (Decapoda, 
Astacidae). Proc. Biol. Soc. Washington 79: 109-116. 

Penn, George H., Jr. 1953. A new burrowing crawfish of the genus 
Procambarus from Louisiana and Mississippi. Tulane Stud. 
Zool. 1: 71-76. 



200 Proceedings of the Biological Society of Washington 



QH 

1 
B4X 



82, pp. 201-204 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



A NEW FLORIDAN CRYPTOPS 

WITH KEY TO THE STATE'S SPECIES^ 

(CHILOPODA : SCOLOPENDROMORPHA : 

CRYPTOPIDAE) 

By R. E. Crabill, Jr. 
Smithsonian Institution, Washington, D. C. 

Heretofore three species of Cryptops (Cryptops) were known 
to inhabit Florida; all were described from that State: hyalinus 
Say,"* floridanus Chamberlin/ and denmarki Chamberlin.* In 
light of what is known of the sporadically widespread dispersal 
of the well-known European hortensis Leach, I feel confident 
that it also occurs in Florida, even though its presence there 
is yet to be detected. 

Through the most recent kindness of Professor E. O. Wilson 
of Harvard University I have been able to secure insular speci- 
mens of a fifth Floridan species, which, believing to be new, 
I describe here. The appended key distinguishes the four spe- 
cies known so far from Florida plus the postulated hortensis.^ 

The new species, parydrus, seems most to resemble the 
Brazilian heathi Chamberlin,** but differs from it as follows. 
In heathi: (1) color yellowish. (2) Paramedian sutures of first 
tergite posteriorly complete. (3) Ultimate leg prefemur and 
femur equal in length. In parydrus: ( 1 ) color distinctly olive 
green. (2) Paramedian sutures of first tergite posteriorly widely 

1 Undertaken with the aid of a grant from the National Science Foundation. 

2Joum. Acad. Nat. Sci. Philadelphia, 2, p. 104, 1821. The best composite de- 
scription of hyalinus is that of Attems in Das Tierreich, 54, p. 224, 1930, but it is 
not without error. Because Attems' figure (number 287) of the cervical con- 
figuration is grossly inaccurate, I present here in my figure 1 a detailed drawing of 
the anterior end of the animal. 

3 Proc. Biol. Soc. Washington, 28, p. 26, 1925. 

* Ent. News, 69, p. 13, 1958. 

5 Having never seen specimens of floridanus and denmarki, which are known 
only from the types, I have had to rely solely upon what information Chamberlin 
presented in their original descriptions. If this information is accurEite, then 
denmarki, floridanus, hyalinus, hortensis and parydrus surely all seem to be 
different species. ^__ ^ . _^. 2>- 

8 BuU. Mus. Comp. Zool. Harvard, 58, p. 157. *** • * ^m^ittm 

15— Proc. Biol. Soc. Wash., Vol. 82, 1969 _^ ( 2Q1J 

JUN26|g69 



202 Proceedings of the Biological Society of Washington 

incomplete. (3) Ultimate leg prefemur notably shorter than 
femur. 

Known Floridan Cryptops 

la. First tergite overlapping rear margin of head. First tergite without 
cervical suture or other sutures hortensis Leach 

lb. Anterior margin of first tergite (in unextended specimens, hence 
normally) overlapped by rear margin of head. First tergite with or 
without cervical and other sutures 2 

2a. First tergite without cervical suture; sutural configuration con- 
sisting of two complete paramedian sutures 

denmarki Chamberhn 



2b. First tergite with cervical suture; sutural configuration not as 
above 3 

3a. Head with long anterior and posterior sutures (fig. 1, A). First 
tergite configuration not omegoid (fig. 1, C). Paramedial sutures 
of 2nd tergite incomplete (fig. 1, H) _.— — hyalinus Chamberlin 

3b. Head with only very short posterior sutural fragments (fig. 2, A). 
First tergite sutural configuration omegoid (fig. 2, C). Paramedian 
sutures of second tergite complete ( fig. 2, H ) 4 

4a. First tergite sutural configuration with complete paramedian su- 
tures. Prostemum medially diastemate floridanus Chamberlin 

4b. First tergite without complete paramedian sutures. Prostemum 
medially not diastemate parydrus new species 

Cryptaps parydrus new species 

Holotype: female. Florida, Dade County, Florida Keys, Bay Point. 29 
June 1967. R. Silberglied, leg. Under bark of red mangrove; taken with 
young. Deposited in the Museum of Comparative Zoology, Harvard 
University. 

GENERAL. Length, 17 mm. Color, distinctly ohve-green. Shape: an- 
terior third of body shghtly attenuate. ANTENNAE. Articles submonih- 
form. Setae distally increasing in number and decreasing in length. 
Length to head length, 9 : 3. HEAD. Slightly longer than wide; shghtly 
wider anteriorly than posteriorly. Paramedian sutures absent, represented 
only by posterior fragments. LABRUM. With a single median tooth (the 
midpiece); sidepieces not incised to form lateral teeth. PREHENSORS. 
Flexed, not exceeding anterior margin of head. PROSTERNUM. An- 
terior margin straight, without diastema, laterally not bowed forward. 
TERGITES. First: cervical suture present; sutural configuration anteri- 



-> 

Fig. 1. Cryptops hyalinus Say, head and first three tergites. A = 
anterior and posterior cephalic sutures. B = cervical suture. C = first 
tergite sutural configuration. D = intertergites. E = obhque suture. 



New Florida centipede 



203 




F = lateral suture. G == paramedian suture. H = posterior fragments of 
paramedian suture. 

Figs. 2 and 3. Cryptops parydrus, holotype, 2. head and first three 
tergites. A = posterior fragment of cephalic suture. B = cervical suture. 
C = first tergite sutural configuration. D =: intertergite. E = obhque 
suture. F = lateral suture. G = paramedian suture. H = complete para- 
median suture of second tergite. 



SMITHSONIAN 
IKSinUTiON 



MAY29196S 



204 Proceedings of the Biological Society of Washington 



orly omegoid, its paramedial sutures broadly incomplete; without oblique, 
lateral, or posterotransverse sutures. Oblique sutures present on 2 through 
18. Complete paramedian and lateral sutures present on 2 through 20. 
STERNITES. Metastemites ( endosternites ) not demarcated by sutures. 
Transverse apodemes present on 2 through 20. All with numerous mi- 
cropores, these scattered, not in formed fields. LEGS. Tarsi 1 through 19 
entire, without trace of division; 20 and 21 completely divided in two. 
Pretarsal parungues on 1 through 20 minute, equal; absent on 21. Spurs 
and spines absent on 1 through 20. ULTIMATE PEDAL SEGMENT. 
Abruptly narrower and shorter than penult. Tergite posteriorly broadly 
pointed. Stemite wider than long, posteriorly broadly, evenly rounded. 
Coxopleural pores few (16-18), on lower fourth of coxopleuron, not 
reaching posterior margin. ULTIMATE LEGS. Relatively short and ro- 
bust. Prefemur notably shorter than femur, which is the longest article. 
Shallowly dorsally sulcate are: distal end of prefemur, entire femur, 
proximal end of tibia. All articles flattened, tibia and tarsalia strongly so. 
Robust spiniform setae present: on prefemur and femur mesally and 
ventraUy; absent on tibia and tarsalia. Mucrones.^ 1 on right femur, 2 
on left (anomalous condition); 6 on right bitia, 5 on left; 2 on each 1st 
tarsus; absent on 2nd tarsus. Ventral surface of 2nd tarsus compressed 
to form a thin, knifelike edge. Pretarsal parungues absent. 

The paratypes agree very closely with the holotype. Apart from one 
notable sexual departure, the chief differences are in length and, occa- 
sionally, in ultimate leg mucro armature. Like the holotype, all are 
Floridan. 

Paratype A: female. Dade County, Florida, Keys, Snipe Key. 27 June 
1967. R. Silberglied, leg. Under bark of red mangrove tree with brood 
of young. Deposited in the U. S. National Museum. Length, 19 mm. 
Mucrones: femora, 1, 1; tibiae, 5, 5; 1st tarsi, 2, 2. 

Paratype B: female. Gilchrist County, south of Trenton. 2 April 1959. 
R. E. Woodruff, leg. Deposited in the U. S. National Museum. Length, 

18 mm. Mucrones: femora, 1, 1; tibiae, 5, 4; tarsi, 2, 2. 

Paratype C: male. Monroe County, Florida Keys, Upper Snipe Key. 
1 August 1967. R. Silberglied, leg. Under red mangrove bark. Deposited 
in the Museum of Comparative Zoology, Harvard University. Length, 

19 mm. Mucrones: femora, 1, 1; tibiae, 5, 5; 1st tarsi, 2, 2. In this single 
male of the series the penult ( 20th ) legs differ conspicuously from those 
of the female. In the male they are considerably swollen, especially ven- 
trally where they are in addition finely and densely setose. In the females 
the penult legs do not differ substantially from the predecents. 



"^ Singular, mucro: an ankylosed spurlike structure occurring ventrally on the 
cryptopid tibia and tarsus. The nomenclature of the different spines and setae and 
their derivatives, as they occur in the Cryptopidae, is discussed in Grabill, Proc. 
U. S. Nat. Mus., Ill (3422), pp. 8, 13-14, 1960. 



^^ 



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B4X 1. 82, pp. 205-208 29 May, 1969 



NH 



PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 

TENONIA KITSAPENSIS, A NEW GENUS AND SPECIES 

OF THE FAMILY POLYNOIDAE (POLYCHAETA) 

FROM PUGET SOUND (WASHINGTON)^ 

•By Frederic H. Nichols 

Department of Oceanography, University of Washington, 

Seattle 

Among the polychaetes collected in grab samples from sev- 
eral stations during studies of the benthic fauna of Puget Soimd 
(Lie, 1968; Nichols, 1968) were many specimens of a new 
species belonging to a new genus in the family Polynoidae 
(listed as Polynoidae sp. I by Lie and Nichols). 

Tenonia new genus 

Type species: T. kitsapensis new species. Gender: feminine. 

Diagnosis: Polynoid with a short body composed of up to 33 segments, 
with up to 15 pairs of elytra. Prostomial lobes rounded without peaks. 
The median antenna inserted dorsoanteriorly; the lateral antennae in- 
serted ventrally. Notosetae and neurosetae numerous; all setae of ap- 
proximately the same thickness. Notosetae and upper neurosetae with 
capillary tips; lower neurosetae with bifid tips. 

Remarks: Of existing polynoid genera, Tenonia most closely resembles 
Antinoe, Gattyana, and Hesperonoe in general body form and type of 
some of the setae. These differ from Tenonia as follows: Antinoe, Kin- 
berg (1855) as redefined by Hartman (1948) has distinct prostomial 
peaks, blunt notosetae, and all neurosetae end in fine tips. Gattyana 
Mcintosh (1900), as emended by Pettibone (1953), has only stout 
neurosetae with simple hooked tips. Hesperonoe Chamberlain ( 1919 ) has 
prostomial peaks, some stout, blunt notosetae in addition to capillary 
notosetae, and subacicular neurosetae with smooth, unidentate tips. 

The name was arbitrarily derived. 

Tenonia kitsapensis new species 
Fig. lA-J 
Material examined: The species description is based on 51 specimens. 
The holotype and 14 paratypes are deposited in the U. S. National 



1 Contribution No. 469 from the Department of Oceanography, University, Qf 
Washington, Seattle, Washington 98105. This study was supported in part 'by' ** '"^"^•"~^'— «.^. 
grant AT (45-1) -1725 from the Atomic Energy Commission, Ref. RLO-1725-120. 

16— Proc. Biol. Soc. Wash., Vol. 82, 1969 (205) 

^THS0/V7^^ 



206 Proceedings of the Biological Society of Washington 




Fig. 1. Tenonia kitsapensis new species: A, dorsal view of anterior 
end; B, left elytron from middle of the body; C, elytral microtubercle; 
D, anterior view of 11th parapodium; E, upper notoseta; F, middle 
notoseta; proximal one-third of serrated blade shown; G, upper neuro- 
seta, proximal one-third of serrated blade shown; H, middle neuroseta; 
I, lower neuroseta; J, cutting edge of lower neuroseta. 



Museum. The holotype (USNM 38263) was collected in Port Madison, 
Puget Sound, Washington, 47°44'08"N, 122°32'00"W, 28 m depth, in 
fine sand with shell fragments (22 October 1965). Of the 14 paratypes, 
also collected in Port Madison, 4 specimens (USNM 38264) were col- 
lected with the holotype; 5 specimens (USNM 38265) at 47°44'09"N, 
122°32'03"W, 30 m depth, in fine sand with shell fragments (21 October 



New Washington polychaete 207 

1965); 4 specimens (USNM 38267, 9 January 1963) and 1 specimen 
(USNM 38266, 29 April 1963) at 47M4'31"N, 122°32'41"W, 12 m 
depth, in very fine sand. Of the remaining 36 specimens ( author's coll. ) , 
30 were collected at various stations in Port Madison along a transect 
between 47°44'04"N, 122°31'54"W and 47°44'35"N, 122°32'44"W, 
2 to 34 m depth, in fine, clean sand to very fine, muddy sand ( 1963 and 
1965); 5 specimens in Case Inlet, southern Puget Sound, Washington, 
47°13'30"N, 122°49'36"W, 68 m depth, in soft mud (1963); 1 speci- 
men near the eastern shore of central Puget Sound, 47°41'33"N, 
122°24'18"W, 23 m depth, in fine gravelly sand (1963). 

The species name is derived from the name of the land area adjacent 
to the type locality, the Kitsap Peninsula. 

Description: Length 3 to 8 mm; width 1.4 to 3.2 mm without setae. 
Segments number 23 to 33. The body shape is oval, widest anteriorly, 
tapering gradually posteriorly. The anterior portion of the body is con- 
vex dorsally, while the posterior is flattened. The body surface is smooth 
and generally transparent. Coloration in formalin is variable, from speci- 
mens totally lacking in color except for the amber color of the pharynx 
seen through the body wall, to specimens with Hghtly speckled body and 
elytra, sometimes with dark intersegmental grooves. 

The prostomium is strongly bilobed, but rounded anteriorly without 
cephalic peaks ( Fig. 1 A ) . Two pairs of large eyes, not seen on many pre- 
served specimens, are positioned in the form of a square as seen from 
above. The anterior pair is located on the anteroventral edge of the 
prostomial lobes but is visible through the transparent prostomium. The 
posterior pair is located dorsally near the posterior edge of the prosto- 
mium. The median antenna with a large ceratophore is inserted antero- 
dorsally in the notch between the prostomial lobes, the long smooth 
style projecting dorsally. The lateral antennae are inserted ventrally on 
distinct ceratophores with smooth bulbous styles approximately one- 
third the length of the style of the median antenna. The palps are smooth, 
thick, tapered, and somewhat longer than the median antenna. The 
tentacular cirri are also smooth and are similar in shape and length to 
the median antenna. The proboscis is well developed, cyhndrical, with 
two pairs of jaws and a circlet of 18 terminal papillae. 

Elytra number 15 pairs on the largest specimen, and are located on 
segments 2, 4, 5, 7, 9, . . ., 21, 23, 26, 29, and 32. There are fewer 
elytra on all other specimens, however, and in some cases there are only 
13 or 14 pairs of elytra (29 to 31 segments) on what appear to be nearly 
mature females. On the first eight to ten segments the middorsum is not 
covered by the elytra. The elytra are pale and smooth with the excep- 
tion of a clearly marked system of veins and a few microtubercles on the 
midlateroposterior region (Fig. IB, C). Elytral margins are smooth and 
transparent. The perimeter of the elytral scar is marked with a circle of 
brown pigmentation on some of the specimens. 

Parapodia are biramous with long smooth dorsal cirri, short pointed 
notopodia, long neuropodia, and ventral cirri of normal size (Fig. ID). 






208 Proceedings of the Biological Society of Washington 

The ventral cirrus of the first setiger is as long as the parapodium. Acicu- 
lae protrude sHghtly from the neuropodial lobes. 

All setae are of approximately the the same thickness ( 5 to 10 /* ) and 
are characteristically long, thin, and colorless (Fig. lE-J), with longi- 
tudinal striations seen under high magnification. All notosetae are capil- 
laries and are serrated for about one-third to one-half of their lengths 
(Fig. IE, F). Upper notosetae (Fig. IE) are short and bent. Lower 
notosetae are long and straight. Upped neurosetae (Fig. IG) end in 
capillary tips, thereby resembling the notosetae. Lower neurosetae (Fig. 
IH, I, J) are bifid. There are about 30 to 40 notosetae and 40 to 60 
neurosetae in the largest specimen. 

The pygidium is blunt with two long smooth anal cirri similar in size 
and shape to the dorsal cirri. 

Diagnosis: A Tenonia species with smooth elytra, except for veins and 
few microtubercles. Elytra do not cover the middorsum of anterior 
segments. Eyes large, forming a square when viewed from above; the 
anterior pair located on the anteroventral edge of the prostomium; the 
posterior pair near the posterior edge. All setae slender, colorless, with 
serrated edges. Palps, antennae, and cirri smooth. 

Distribution: Puget Sound, Washington; 2 to 70 meters, in sand and 
sandy mud. There were about 15 specimens/m^ at the type locality. 

Literature Cited 

Chamberlin, R. V. 1919. Pacific coast Polychaeta collected by Alex- 
ander Agassiz. Bull. Mus. Comp. Zool. Harvard, 63(6) :251- 
270; pis. 1-3. 

Hartman, O. 1948. The marine annelids erected by Kinberg with 
notes on some other types in the Swedish State Museum. 
Ark. Zool. Stockholm, 42A(1): 1-137; pis. 1-18. 

Kinberg, J. G. H. 1855. Nya slagten och arter af AnneHder. Ofv. Vet. 
Akad. Stockholm, Forh. 12(9):381-388. 

Lie, U. 1968. A quantitative study of benthic infauna in Puget Sound, 
Washington, U.S.A., in 1963-1964. FiskDir. Skr. Ser. 
Haviinders. 14(5): 229-556. 

McIntosh, W. C. 1900. A monograph of the British annelids. XL 
Polychaeta. Amphinomidae to SigaUonidae. Ray Soc. London, 
215-442; pis. 24-42. 

Nichols, F. H. 1968. A quantitative study of benthic polychaete as- 
semblages in Port Madison, Washington. M.S. Thesis, Univ. 
Washington. 78 p. 

Pettibone, M. H. 1953. Some scale-bearing polychaetes of Puget 
Sound and adjacent waters. Univ. Washington Press, Seattle. 
89 p. 



QH 

1 

B4X I. 82, pp. 200-218 29 May, 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



NEW SPECIES OF SIPUNCULA FROM THE 
WESTERN NORTH ATLANTIC 

By Edward B. Cutler 

Biology Department Utica College of Syracuse University, 

Utica, New York 

Among the sipunculans collected during the past few years 
off the east coast of the United States (Emery and Schlee, 1963; 
Sanders, Hessler and Hampson, 1965; Cutler, 1967) were sev- 
eral new species, four of which are herein described. All of 
them were small (less than 1 cm) and came from fairly great 
depths (off the continental shelf). 

This work is part of a larger study (Cutler, 1967) which may 
be referred to for more complete information about the spe- 
cies' distribution. This larger work will be submitted for publi- 
cation in the near future. 

I wish to acknowledge the support of the Cooperative 
Oceanographic Program of Duke University during part of 
this work. 

Aspidosiphon zinni new species 

Diagnosis: A very small Aspidosiphon with a fine-grained, ungrooved 
anal shield from deep water. 

Description: Two hundred and forty six specimens from 17 stations. 

These worms are small (1-5 mm) and fragile. Most are 2-3.5 mm 
long by about 0.3 mm in diameter. They are cylindrical and transparent 
except at the two ends where there is some light brown pigmentation. 
The introvert is almost equal in length to the trunk and expands near 
its tip giving it the appearance of a spiked club (Fig. lA). Commonly, 
these are found in elongate, arenaceous foraminiferan tests. 

The numerous irregularly arranged hooks are single pointed and about 
15-20 fi from base to tip (Fig. IC). No tentacles were observed. The 
anal shield is yellowish and composed of fine, closely-grouped units 
giving it a granular appearance ( Fig. IB ) . This shield is flat and usually 
set off from the main axis of the trunk by 60 to 90 degrees. The posterior 

17— Proc. Biol. Soc. Wash., Vol. 82, 1969 (209) ] 

/^THS0/V7^^ 

26t969 



( JUN2 



210 Proceedings of the Biological Society of Washington 




Figure 1. Showing Aspidosiphon zinni: (A) the entire animal show- 
ing anus (a) and intestine (i) through body wall: (B) the enlarged 
portion of anal shield: (C) an isolated hook from introvert. 



shield is very poorly defined, but annular wrinkles or folds are discern- 
ible at the posterior end of the trunk. 

It is possible to see most of the internal structures through the body 
wall; however, all attempts at dissection have been unsuccessful. Attempts 
were made to section the material but these did not yield any useful 
information other than that the longitudinal muscle layer was continuous. 
The intestinal coil extends to the posterior end of the trunk and the anus 
is just behind the anterior shield. The two brown nephridia open at about 
the level of the anus and are usually less than 25 percent of the trunk 
length. Two retractors are present and originate near the ventral nerve 
cord about 65-80 percent of the distance to the posterior end of the trunk. 

Remarks: The only other member of this genus thus far reported from 
similar latitudes is Aspidosiphon mulleri Diesing, 1851, and with one 
exception (Fischer, 1922:22), it seems to be restricted to the eastern side 
of the Atlantic Ocean. These two species are quite different; A. mulleri 
is generally larger ( 8-15 mm ) with dark brown, more massive, furrowed 
anal shields. No other species approaches this one in terms of morphology 
or distribution. 

This species has been named for Dr. Donald J. Zinn, Professor of 
Zoology, University of Rhode Island. 



New Atlantic sipunculid 211 




Figure 2. Showing Onchnesoma magnibatha. An external view show- 
ing intestinal coil through transparent body wall. 



Distribution: This species is found between 2000 and 3000 m with a 
few at shallower depths (shallowest 1102 m), generally off the mid- 
Atlantic States. The temperatures are cold and the sediments are very 
fine. This is not the typical Aspidosiphon habitat. 

Holotype: U.S.N.M. 38242; Location 39°26' N, 70°38' W; Depth, 2496 
m; Collected 21 August 1964 by Sanders, Hessler and Hampson. 

Paratype: U.S.N.M. 38243; on sHde; Location 39°42' N, 70°39" W; 
Depth, 2086 m; Collected 24 May 1963 by Sanders, Hessler and 
Hampson. 

Onchnesoma magnibatha new species 

Diagnosis: An Onchnesoma with pronounced, continuous posterior 
ridges and thin, transparent skin from deep, cold water. 

Description: Twenty four specimens from 11 stations. 

The trunk varies in length from 2.5-6 mm, and in width from 1-2 mm. 
The shape varies from almost cyUndrical to pear-shaped with the pos- 
terior end bluntly pointed. The anterior M to /2 of the trunk is narrowed 
to a diameter equal to that of the introvert; only the posterior section 
is bulbous (Fig. 2). 

The skin of the central enlarged portion is very thin, fragile, and 
transparent, while that of the anterior narrowed part is opaque, some- 
what rugose, and pale brown-tan. The posterior end has a series of 
longitudinal ridges merging at the apex and has a suggestion of the tan 
coloration. These ridges are more pronounced than in Onchnesoma steen- 
strupi Koren and Danielssen, 1875, and are more continuous i.e., not 
composed of distinctly separate scales or plates. 

The introvert is generally the same diameter as the anterior part of 
the trunk but somewhat paler; it is smooth and transparent if extended. 
None of my material is completely extended so I cannot comment on the 
tentacular arrangement. The introvert appears to be less than twice the 
length of the trunk and often (measured on dissected, partially con- 
tracted specimens) only equal in length to the trunk. 



SMIIHSONIAK MfiYOQiqfiQ 



212 Proceedings of the Biological Society of Washington 



W1/ff 




Figure 3. Showing Golfingia (Mitosiphon) murinae. (A) is an ex- 
ternal view with intestine showing through body wall. This individual 
has posterior papillae which are more prominent than some others: (B) 
shows several nephridia from G. murinae hilohatae, with varying degrees 
of development of second lobe. 



The intestine has about 30 double coils; the esophagus and rectum 
are long; there is one or sometimes two fixing muscles, one attaching 
the esophagus near the posterior end of the body. The anus is located 
about 75-80 percent of the distance towards the distal end of the intro- 
vert. The single nephridium is on the right side, unpigmented, and un- 
attached to the body wall except at the nephridiopore. The retractor 
appears single in most contracted specimens, but in one case, there are 
two distinct roots. 

Remarks: This species superficially resembles some members of the 
genus Golfingia Lankester, 1885, more than the other Onchnesoma. 
There is no question, however, that it does belong to Onchnesoma. It has 
the single retractor, single nephridium, and anteriorly displaced anus. 
Internally there is not much to distinguish this species from the other 
two members of the genus, O. steenstrupi and O. squamatum Koren 
and Danielssen, 1875. It has none of the large scalelike papillae char- 



New Atlantic sipunculid 213 

acteristic of O. squamatum, but does somewhat resemble O. steenstrupi. 
It differs from the latter species in having a much shorter introvert; the 
anus not immediately behind the mouth; a much thinner, transparent 
skin; posterior ridges more pronounced and continuous; a pair of roots 
sometimes present on the retractor; and a somewhat different shape 
and color. 

Distribution: Onchnesoma magnibatha (as the name implies) has 
been collected only from great depths, generally between 3750-4980 
m off the mid-Atlantic States where the temperatures are stable and cold; 

Holotype: U.S.N.M. 38244; Location 33°57' N, 65°47' W; Depth, 
4715 m; collected 1 May 1966 by Sanders et al. 

Paratypes: U.S.N.M. 38245; Location 00°46' S, 29*28' W; Depth 3459 
m; collected 14 February 1967 by Sanders et al. 

Golf ingia murinae new species 

Diagnosis: A Golfingia in the subgenus Mitosiphon with long, slender 
introvert, large mammiform papillae on the posterior end of the trunk 
and hooks with a comblike series of teeth at the base of some of the 
hooks. May have single or bilobed nephridia. 

Description: These worms have trunks 1.5-13 mm. The introvert is 
at least four to six times the length of the body. They are semi-transpar- 
ent, yellow, tan, or brown depending on their size and state of contrac- 
tion. The trunk is generally shaped like a plump spindle with mammi- 
form papillae on the posterior end which are usually very distinct (Fig. 
3A). The anterior delineation of the trunk is not always clear as it often 
tapers gradually into the long, delicate introvert. Small hooks ( 25-40 fi ) 
are present at the tip of the introvert in a few rows, then becoming 
sparsely scattered. These hooks just behind the tentacles have an acces- 
sory comb of spinelets at the base. These are easily overlooked. The nature 
of the tentacles can not be ascertained because no specimen is completely 
expanded. There are small papillae on the remainder of the introvert. 

The trunk wall in some specimens is rather thick and the rough, loose- 
appearing epidermis of some specimens has blister-Hke papillae. Smaller 
ones are thin-skinned and transparent. The papillae are discussed at 
length in Murina (1964b) and measurements are presented in her Table 
3. She divided them into three types: rosette-shaped, dome-shaped with 
crowns, and oval bodies. The most outstanding external feature of this 
species is the large posterior papillae. Some typical measurements are 
(length and breadth in microns) : 85 X 75, 157 X 91, 200 X 114, 227 X 
200. They often measure 55-65 fi in height. These measurements are of 
the same order of magnitude as Murina's oval bodies. 

Internally the four retractors are more typically arranged than in the 
Golfingia trichocephala Sluiter complex i.e., the ventral pair is usually 
larger than the dorsal pair, and both pairs are set off farther from the 
ventral nerve cord, the dorsals more so than the ventrals. The origin of 
the retractors generally lies in the posterior third of the body (65-85% 
of the distance to the posterior end of the trunk) but may be found 



214 Proceedings of the Biological Society of Washington 

farther anterior. The single lobed nephridia are free, sac-shaped, and 
open at the anterior of the trunk. The bilobed nephridia have a secondary, 
anterior lobe of varying dimensions (Fig. 3B). The anus is always pos- 
terior to the nephridiopores but the distance may vary. One and some- 
times two fixing muscles were observed (Murina found up to three). 
The intestine is fixed to the posterior end of the body by the spindle 
muscle. 

Since submission of the manuscript one specimen from recent collec- 
tions was found with its introvert completely extended. The 8-10 very 
small, stubby tentacles are arranged in a cluster off to one side very much 
like the situation in the genus Phascolosoma. The location of the mouth 
could not be confirmed. 

Remarks: This species is closely related to Golfingia hespera but differs 
in four ways. The only external difference is the nature of the papillae. 
These are larger and more apparent on G. murinae and more clustered to- 
wards the posterior end while in G. hespera they are smaller, more scat- 
tered, and sometimes pigmented. Internally the differences are in the 
nephridia and retractor muscles. The nephridia in G. hespera are always 
bilobed, the two lobes being more or less equal, long, thin lobes. In those 
G. murinae which have two lobes they are not of equal size, one usually 
significantly shorter than the other. The retractors in G. murinae are 
more typical with the ventral pair larger and both pairs set off farther 
from the ventral nerve cord with the ventrals being closer to the nerve 
than the dorsals. This is not the case in G. hespera. 

In addition to the morphological differences, there are ecological 
differences which I assume to be real and meaningful. G. hespera has 
been reported from shallow, warm or temperate areas while G. murinae 
comes from deep, cold areas. Their distribution patterns are distinct. 

The following two subspecies may represent a dynamic situation in 
which one form is the ancestral stock, and the other has only recently 
separated from it. This will be investigated in more detail in the future. 

Murina (1964a) has described two Golfingia from 395 m in the 
Northwestern Pacific and several from the Mediterranean ( 1964b ) which 
she placed in the species Golfingia hespera. For reasons discussed else- 
where (Cutler, 1967) it is apparent that Murina's designation was in- 
correct. Her material represents, in my opinion, two subspecies of a new 
species which I have named after her in recognition of her contributions 
to our knowledge of the Sipuncula. 

Golfingia murinae unilobatae new subspecies 

Diagnosis: Two hundred and one specimens from 35 stations. 

This subspecies fits the description of the species but has only the 
single lobed nephridia. It is generally larger than the other subspecies 
(4-13 mm), but the size ranges overlap. This form serves as the type 
for the species. Included in this subspecies would be Murina's ( 1964a ) 
Pacific specimens of G. hespera. 



New Atlantic sipunculid 215 

Holotype: U.S.N.M. 38247; Location, 37°13' N, 68°40' W; Depth, 
4540 m; collected 25 May 1962 by Sanders et al. 

Paratype: U.S.N.M. 38248; Location, 37n3' N, 68M0' W; Depth, 
4540 m; collected 25 May 1962 by Sanders et al. 

Golf ingia murinae bilobatae new subspecies 

Diagnosis: Eighty-eight specimens from 22 stations. 

Morphologically this subspecies is almost identical with the preceding 
one. The trunk size is smaller; these range from 2-7 mm as compared 
with a maximum of 13 mm in the former. As before, the striking feature 
is the presence of large, mammiform papillae on the posterior end of the 
trunk. However, there is one internal difference; a small, secondary 
anterior lobe on the nephridia. If this is absent ( 3 out of 36 ) the nephrid- 
ium is long and slender in contrast to the short, fat one in G. m. uni- 
lobatae. This by itself may not be just cause for erecting a separate 
subspecies if it were not for the distinctive distribution pattern of this 
bilobed form. Murina's ( 1964b ) Mediterranean specimens of G. hespera 
would be included in this subspecies. 

Holotype: U.S.N.M. 38249; Location, 24°10' N, 81°22" W; Depth 
677 m; collected 1 June 1964 by R. Wigley. 

Paratype: U.S.N.M. 38250; Location, 24°10' N, 81°22' W; Depth, 
677 m; collected 1 June 1964 by R. Wigley. 

Distribution: The distribution of these two subspecies is spatially sep- 
arate. G. murinae bilobatae is found on the continental slope from Cape 
Hatteras, North Carolina, south to the Florida Keys (34° 16' N to 24*10' 
N) at depths of about 300-700 m. G. murinae unilobatae is found from 
32° 11' to 39° 59' N. The most southerly ones are all on the Bermuda 
Slope between 1000-2000 m. The remainder are at depths between 2200 
m and 4750 m on the continental rise between Cape Cod, Massachusetts, 
and Cape Hatteras, North Carohna. 

In summary, this species prefers a relatively cold, stable, deep environ- 
ment with fine sediments. One subspecies is found in shallower warmer 
areas with a slightly coarser substratum than the other. 

Golf ingia constricticervix new species 

Diagnosis: A Golfingia in the subgenus Phascoloides with an unusual 
neck like constriction and short introvert with large hooks on the distal 
half from moderately deep, cold water. 

Description: Nine specimens from 4 stations. 

These are long, slender animals, the length being about 15-20 times 
the diameter ( 15 X .67 mm; 5 X .33 mm). They are generally transparent 
becoming dark yellow towards the two ends of the trunk. The introvert 
is about 1/5 the length of the trunk (Fig. 4A). 

One of the most unique features of this species is a constriction or neck 
at the base of the introvert. Because of this relatively weak connection, 
few worms are whole. Those few retaining the introvert show the long 



216 Proceedings of the Biological Society of Washington 




Figure 4. Showing Golfingia (Phascoloides) constricticervix: (A) is 
the entire animal with partially expanded introvert: (B) is completely 
expanded introvert shovidng tentacles. 

hooks which cover its distal half. The hook size increases from 0.04 mm 
to 0.28 mm. This marked increase in size towards the tip has not been 
recorded for other species. The hooks are single pointed, moderately 
curved, slender, and irregularly arranged. About 12-16 short, reduced 
tentacles are visible on one individual (Fig. 4B). 

The anterior end of the trunk is usually swollen and the anus is located 
on a small protruberance. Very small, slightly oval, clear, glandular 
papiUae are visible at the anterior and posterior ends of the trunk which 
is otherwise smooth and thin-walled. 



New Atlantic sipunculid 



217 



Internally the two retractors originate from the body wall very close 
to the ventral nerve cord about 15-25 percent of the distance to the 
posterior end of the trunk. These two muscles join together before leaving 
the trunk as one unit. The intestinal coil may be loose and the individual 
coils are often located relatively far apart from one another. 

Remarks: This species is similar to Golfingia vitjazi Murina, but lacks 
the round, ribbed, chitinous shield of the anterior end of the trunk. 
Specimens of G. constricticervix were sent to Dr. Murina and she agreed 
that they were two different species. 

These worms are easily identified when the introvert is extended and 
attached, but if it is withdrawn or broken off, it has a non-descript, non- 
sipunculan appearance. The specific name refers to the outstanding 
morphological feature of a constricted "neck" region. 

Distribution: This species was found near Bermuda between 1700 m 
and 4800 m and further north on the continental rise and slope, generally 
between 1500-3000 m. The temperatures are cold and the sediments 
quite fine. 

Holotype: U.S.N.M. 38246; Location, 33°57' N, 65°47' W; Depth, 
4815 m; collected 1 May 1966 by Sanders et al. 



Literature Cited 

Cutler, E. B. 1967. The Sipuncula from the Western North Atlantic: 
their systematics, ecology, and distribution. Unpublished 
doctoral dissertation. University of Rhode Island. 258 p. 

Emery, K. O. and J. S. Schlee. 1963. The Atlantic Continental Shelf 
and Slope, a program for study. Geol. Survey Circ. 481 U.S. 
Dep. of Interior. 11 p. 

Fischer, W. 1922. Gephyreen des Reichsmuseums zu Stockholm 
Arkiv for Zoologi. Band 14, no. 19: 1-39. 

Murina, V. V. 1964a. New and rare species of sipuncuUds of genus 
Golfingia. [In Russian] Works Inst. Oceanol. Acad. Sci. 
USSR 64: 216-253. 

. 1964b. Sipuncuhd fauna of the Mediterranean Sea. [In 

Russian] Works Sevastopol Biol. Sta. 17: 51-76. 

Sanders, H. L., R. R. Hessler, and G. R. Hampson. 1965. An in- 
troduction to the study of deep-sea benthic faunal assem- 
blages along the Gay Head-Bermuda transect. Deep-Sea Res. 
12: 845-867. 



218 Proceedings of the Biological Society of Washington 



/.■' 



^^V.• 



82, pp. 219-232 29 May, 1969 

PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



TWO ADDITIONAL SUBSPECIES OF NORTH 

AMERICAN CROTALID SNAKES, 

GENUS AGKISTRODON 

By Howard K. Gloyd 

Department of Biological Sciences, 

University of Arizona, Tucson, Arizona 

Investigations leading toward a monographic review of the 
genus Agkistrodon Beauvois, 1799, have indicated that the 
recognition of some additional subspecies will tend to make 
the pattern of geographic dispersal and probable evolu- 
tionary history of the group more easily interpreted. In this 
paper a fifth subspecies of the copperhead, A. contortrix ( Lin- 
naeus), and a third subspecies of the cottonmouth, A. pisciv- 
orus (Lacepede), are described. In outlining these subspecies, 
a large number of specimens has been examined in each case, 
but since it is not feasible to deal here with the entire hypo- 
digm, diagnoses and definitions are based on a small series 
selected from the central portion of the geographic range of 
each. 

It has been pointed out (Gloyd and Conant, 1943, p. 147) 
that both the copperhead and the cottonmouth, as species, are 
exceedingly homogeneous in structural characters but that 
geographic races are recognizable on the basis of differences 
in color and pattern. Such differences, however, are often sup- 
ported by average ( mean ) differences in at least some morpho- 
logical attributes. The several subspecies now recognized are 
easily distinguished from each other throughout most of their 
respective ranges, but there are many areas in which individ- 
uals are obviously intermediate; inter gradation appears to 
occur wherever any two adjacent subspecies are in contact. 
The keys to the subspecies included here are based upon all 

18— Proc. Biol. Soc. Wash., Vol. 82, 1969 (219) 



(JUK 26 1969) 



220 Proceedings of the Biological Society of Washington 

data presently available to the writer and, along with consid- 
eration of the geographic ranges given, should identify indi- 
viduals other than those definitely intermediate; i. e., at least 
75 percent in each of the respective populations. Discussion 
of geographic and evolutionary relationships is reserved for 
a detailed treatment of the genus. Names of museums abbrevi- 
ated in the text are listed on page 231. 

Agkistrodon ccntortrix phaeogaster new subspecies 
OSAGE COPPERHEAD 

Figure 1 

Agkistrodon mokeson mokeson, Gloyd and Conant, 1943, Chicago Acad. 

Sci. Bull., vol. 7, no. 2, p. 150 (part). 
Agkistrodon contortrix mokeson, Klauber, 1948, Copeia, no. 1, p. 8 (part). 

Holotype: United States National Museum no. 165955, adult male, 
received from Dr. Henry S. Fitch, July, 1960; locality 10 miles south o£ 
McLouth, Jefferson County, Kansas; collector not stated. 

Paratypes: The following, all from east-central Kansas: Douglas County, 
USNM 165958-9, 7 mi. SE of Lawrence; AMNH 102614-5, FMNH 
178997-8, UMMZ 128842, 3 mi. NW of Lawrence, UCM 36348, 7 mi. 
SW of Clinton. Franklin County, USNM 165956-7, Gould's Ford, Middle 
Creek, 3 mi. S and 3 mi. E of Ottawa. Jefferson County, KU 120336-7, 
12 mi. N of Lawrence. Johnson County, AMNH 102616, UMMZ 128843, 
2 mi. S of Stanley. Miami County, USNM 165960-1, Miami County 
State Lake. All specimens here mentioned were collected by Dr. Henry 
S. Fitch and his associates at the University of Kansas, particularly 
Robert Fleet, Richard Hayworth, William Kerfoot, Roy and Dennis 
O'Connor, and Eric Schulenberger, 1960-63. 

Diagnosis: A population of copperheads in which the crossbands of 
the body are ( 1 ) generally in sharp contrast with the ground color; ( 2 ) 
have dark edges narrowly bordered with white, extending down the 
sides to, or almost to, the ventrals; and (3) are 3 to 5 scales wide at 
the midline and 6 to 11 scales wide where they are in contact, or nearly 
so, with the ventrals ( Fig. 1 ) . There are usually no small dark spots 
or blotches in the ground color between the crossbands. The ventral 
markings are irregular gray to black blotches, mostly in contact, extend- 
ing across the belly and producing a dusky, marbled or clouded pattern, 
sometimes quite dark. The yellow tip of the tail of juveniles is replaced 
by yellowish green and this color is retained by most adults. 

Description of Holotype: Top of head with nine symmetrical scutes 
characteristic of the genus, with the frontal aberrant: a longitudinal suture 
divides it, separating on the right side a narrow extra scale about one- 
fourth the total width of the frontal; also a transverse suture separates 
a minute scale at the posterior midpoint of the frontal. Each parietal has 
a dark spot of pigment near its mesial border and back of each of these a 



Two new snake subspecies 



221 




Fig. 1. Agkistrodon contortrix phaeogaster new subspecies. A para- 
type, USNM 165957, Gould's Ford, Franklin County, Kansas. 



SWilHSONIAN MAY291969 



222 Proceedings of the Biological Society of Washington 

short groove extends anterolaterally for about one-third the width of 
the scute. Nasals two, the anterior one larger. Loreals one on each side, 
large, higher than wide. Preoculars two, the lower one forming the upper 
posterior border of the pit. Postoculars (including suboculars) 5 on the 
left side, 4 on the right. A small postfoveal between lower preoculars 
and anteriormost postoculars, making a total of 9 scales in the orbit on 
the left side and 8 on the right. Supralabials 8-7, the second forming 
the anterior border of the pit, the fourth largest and separated from the 
orbit by the two anteriormost post (sub-) oculars. An anterior postfoveal 
forms the lower posterior border of the pit. Infralabials 10 on the left 
side, 9 on the right. Temporals in 6 or 7 irregular rows; the large scales 
of the lowest row, 4 on the left and 3 on the right, conspicuously larger 
than all the others and without keels. The first pair of infralabials in 
broad contact behind the mental. One pair of enlarged chin shields. 
Median gulars in 3 irregular pairs; lateral gulars in 5 to 6 oblique rows. 
Dorsal scales all strongly carinate on body and tail, and each with two 
apical pits. Scale rows 27-25-23-21, the order of decrease (Dowling 
recount system, 1951) as follows: 

-7(15) -5(22) -5(123) 

27 25 23 21 (148) 

-7(16) -5(22) -4(103) 

Ventrals 148; anal plate not divided. Both hemipenes everted. Sub- 
caudals 46, the proximal one and the distal 16 divided. There is a vertical 
scar from a healed wound on the right side of the body on scale rows 4 
to 8 opposite ventral 87, and an indication of a slight injury on the top 
of the snout at the meeting point of the internasals and prefrontals. 

Total length 990 mm; tail length 125 mm; tail/total length ratio .126. 

General coloration in life rich tones of brown: reddish brown on top 
of the head and in the crossbands of the body, with the ground color 
a lighter tone of brown; after preservation faded to corresponding tones 
of darker and lighter grayish brown. Muzzle and top of head uniform, 
with no trace of pattern; a darker tone of brown extending down over the 
temporals forms an indistinct cheek stripe which is outlined below by a 
narrow dark brown line edged with cream color; this extends obliquely 
backward from the orbit across the lowest row of temporals and down- 
ward across the last supralabial to the commissure. At the sides of the 
muzzle above the nares a narrow dark line on the edges of the inter- 
nasals and nasals. Supralabials, lower portions of first row of temporals, 
and upper halves of infralabials cream color, densely stippled with 
brown. A sharp narrow line of dark brown extends backward from the 
second infralabial across the middle of the infralabials to a point just 
below the angle of the mouth; here it curves downward and outlines the 
pale tan pattern of the lower jaw and throat; this area is slightly darker 
than the ventral ground color and is densely stippled with fine dots of 
brown and gray. 

Dorsal ground color grayish brown, with much fine stippling of 



Tivo new snake subspecies 223 

brown and gray. A pattern of dark brown crossbands, all extending down 
the sides and making contact with the ventral scutes. There is a half- 
band on the left side between the first and second crossbands, the next 
nine crossbands are complete, then follow two half-bands on the left 
and one on the right, then two irregular complete bands, a half-band 
on the right, and a final band complete but irregular. Thus a count of 
half-bands on the sides gives 16 on the left and 15 on the right. The 
crossbands vary from 5 to 8 scales wide at the base and 3 to 5 scales 
wide at the midline; they are darker at the borders and narrowly edged 
with light cream (or white). There are no dark spots or blotches in the 
ground color between the crossbands. The tail has one complete band 
proximally, then an irregular zig-zag pattern followed by a dark terminal 
spine. 

Ventral ground color cream to pale tan with fine dark stippling 
throughout the length of the body. A ventral pattern of broad dark 
brown blotches, confluent mesially and extending irregularly toward 
the sides where the outer portions are darker, the whole giving a dusky, 
marbled effect, darker anteriorly and becoming progressively lighter to- 
ward the tail. Under surface of the tail with dense dark stippling, distally 
becoming almost black. 

Notes on Paratypes: Minor variations from the usual scutellation of 
the head include irregular sutures in the parietals extending laterally from 
the mesial plane part way across the scutes. The parietals are usually 
truncate posteriorly. The frontal often with small scales separated off 
the anterior and/or posterior midpoints, or off the anterolateral angles 
(corners). Supralabials usually 8, occasionally 7; infralabials 9 or 10, oc- 
casionally 11. Preoculars 2, a small postfoveal usually entering the orbit 
below them. Postoculars (including suboculars) 4 or 5. Ventrals 143-149 
(146) in 7 males, 143-150 (146) in 9 females. Subcaudals 45-47 (45.7) 
in males, 39-46 (43.1) in females. Divided subcaudals 11-22 (males), 
10-28 (females). Tail/total length ratio .137-.155 (.146) in males, .129- 
.147 (.139) in females. 

In the 16 specimens of the paratype series the crossbands of the body 
range from 12 to 14; in two the 14 crossbands are complete and sym- 
metrically continuous across the body; in the others there are irregulari- 
ties of incomplete bands in which from one to several half -bands alter- 
nate or make only partial contact at the middorsal line. On the tail 
are 1 to 4 proximal crossbands similar to those on the body, followed 
by a dark zig-zag pattern or by a series of 8 or 9 crossbands decreasing 
in size and distance apart distally; terminal spine with 1 or 2 transverse 
black bands. 

In ten of these specimens, medium-sized adults, a lyre-shaped pattern 
on the top of the head and the sides of the neck is more or less clear. 
This is a forward continuation of each of the lateral points of the first 
crossband (on the neck) to become confluent with the dark cheek stripe 
extending forward to the orbit, and expanding mesially into rounded ex- 
tensions two to four scale-widths back of the parietals. Such a pattern 



224 Proceedings of the Biological Society of Washington 

is not evident in large adults in which the entire top of the head, includ- 
ing the postparietal region, is uniform copper-brown. 

The ventral coloration is similar to that described for the holotype 
except for variation in intensity of pigmentation; the belly pattern of 
USNM 165957 (Fig. 1) and USNM 165961 is considerably darker, ap- 
pearing brownish black anteriorly. 

The name phaeogaster alludes to the dusky, dark gray to black, mottl- 
ing of the belly. 

Geographical distribution: A. c. phaeogaster occupies the major portion 
of the Marais des Cygnes-Osage River drainage system in eastern Kansas 
and central Missouri. It inter grades with A. c. laticinctus in southern 
Kansas and northern Oklahoma, and with A. c. contortrix from southern 
Missouri northward along the Mississippi River to southeastern Iowa. 

Key to the Subspecies of the Copperhead 

1. Dorsal pattern of the body consisting of broad dark crossbands, not 
much narrower middorsally than on the sides (more than half as 
wide at the midline as on the lowest lateral scale rows), extending 
across the lateral ends of the ventral scutes and more or less con- 
tinuous with the dark markings of the belly 2 

Dorsal pattern of the body consisting of dark crossbands (or alter- 
nating half -bands) considerably narrower middorsally than on the 
sides (usually less than half as wide at the midline as on the lower 
lateral scale rows), and not continuous with the dark markings of 
the belly 3 

2. Belly very dark, heavily mottled, sometimes almost uniform black; 
crossbands with three dark ventrolateral areas each continuous with 
the dark color of the belly (Fig. 2); tail relatively long, about 17 
percent of total length in males, 14 percent in females; subcaudals 

48-62 (54) in males, 42-57 (49) in females 

pictigaster Gloyd and Conant, 1943. 



Trans-Pecos Texas (Jeff Davis, Presidio, Brewster, and Terrell 
counties) and adjacent areas to the east (Crockett and Val Verde 
counties ) . 

Belly not conspicuously dark; crossbands with three dark ventro- 
lateral blotches extending onto the ventral scutes but tending to 
become weak toward the middle of the belly (Fig. 3); tail about 
14 percent of total length in males, 13 percent in females; sub- 
caudals 42-54 (47) in males, 40-51 (45) in females 

laticinctus Gloyd and Conant, 1934. 

South-central Texas (Frio and Calhoun counties) northward 
through central Oklahoma to southern Kansas (Cowley County). 
Crossbands extending down the sides to the ventral scutes or to 
the lowermost row of dorsal scales, but not continuous with the dark 
irregular blotches of the belly, and without three ventrolateral 
blotches or spots within or immediately adjacent to them ( Fig. 1 ) ; 
belly pattern irregularly mottled or marbled, more dusky ante- 



Two new snake subspecies 



225 







^^0^- 





Fics, 2-5. 2, A. c. pictigaster, a paratype, CA 7807, Oak Spring, 
Chisos Mountains, Brewster County, Texas. 3, A. c. laticinctus, holotype, 
UMMZ 75599, 26 mi. NW of San Antonio, Bexar County, Texas. 4, A. c. 
mokasen, USNM 166134, Scioto County, Ohio. 5, A. c. contortrix, CA 
5089, Gentilly, Orleans Parish, Louisiana. 



226 Proceedings of the Biological Society of Washington 



riorly; no small dark spots in dorsal ground color between cross- 
bands phaeogaster, new subspecies. 

See Geographic distribution above. 

Crossbands not extending down the sides to the ventral scutes, but 

fading out or rounding off on scale row two or above 4 

4. Crossbands somewhat "dumbbell-shaped," moderately restricted at 
the middorsal line (here usually 3 to 5 scale-lengths in width); 
a series of dark ventrolateral spots on the lowermost dorsal scale 
rows and the edges of the ventral scutes, one opposite the middle 
of each crossband and the adjacent ones between the crossbands, 
and all of about equal intensity; usually a secondary pattern of 
small to medium-sized dark spots dorsally between the crossbands, 
sometimes singly, often in pairs ( Fig. 4 ) .___ mokasen* Beauvois, 1799. 
Southern Illinois and northeastern Mississippi northeastward to 
Massachusetts; throughout the Appalachian mountain region, and 
intergrading with contortrix across a broad area of the piedmont 
and Atlantic Coastal Plain. An upland form. 

Crossbands somewhat "hourglass-shaped"; greatly restricted at the 
middorsal line (here 2 or 3 scale-lengths in width), often irregu- 
larly divided, the half-bands not meeting middorsally but alter- 
nating on opposite sides of the body; a series of ventrolateral spots 
not of equal intensity, those opposite the middles of the crossbands 
pale and diffuse, those alternating between the crossbands dark and 
well-defined; no secondary pattern of small dark spots dorsally be- 
tween the crossbands (Fig. 5 ) ; both ground color and pattern gen- 
erally pale contortrix Linnaeus, 1766. 

Mississippi Valley, Culf and Atlantic Coastal Plains, from south- 
eastern Texas north to east-central Missouri and western Illinois, 
and to southern Maryland in the east, exclusive of peninsular 
Florida, A lowland form. 

Agkistrodon piscivorus conanti new subspecies 

FLORIDA COTTONMOUTH 

Figure 6 

Agkistrodon piscivorus piscivorus, Gloyd and Conant, 1943, Chicago 
Acad. Sci. Bull, vol. 7, no. 2, p. 166 (part), fig. 6, 14, and 16. 
Holotype: United States National Museum no. 165962, a young adult 
male, collected 16 July 1966, "at edge of Rochelle-Cross Creek Road, 
about 7 miles southeast" of Gainesville, Alachua County, Florida, by 
R. P. Elliott, J. Wariner, and P. Pinnel. 

Paratypes: USNM 165963, AMNH 102618, Paynes Prairie, about 5.5 
mi. S of Gainesville, Alachua County; USNM 165964, Hampton, Brad- 
ford County; FMNH 178999, 5 mi. E of Andalusia, Flagler County; 
AMNH 102617, UCM 36350, Waccasassa River, 0.5 mi. S of State Road 
24, Levy County; and FMNH 179000, 5 mi. NE of Palatka, Putnam 



'" For explanation of return to this spelling, see Smith and Gloyd, 1964. 



Two new snake subspecies 



227 



' '•; '4"- ^ . 




Fig. 6. Agkistrodon piscivorus conanti new subspecies. CA 11346, 
Florida, exact locality unknown. 

County, Florida. The specimens in this series were collected by Dr. W. J. 
Riemer, J. A. Holman, Norm Tessman, and W. G. Weaver, Jr. of the 
University of Florida. 

Diagnosis: The distinguishing characters of this subspecies most readily 
utilizable are the strong, sharply defined markings of the snout and head, 
even in most relatively large (old) individuals: a dark brown cheek 
stripe outlined above and below by a distinct light line ( white or cream ) ; 
a conspicuous vertical dark stripe on the sides of the rostral and the 
adjacent prenasals and supralabials (Fig. 6); dark markings on labials 
above and below the commissure, more numerous below; and a pair of 
longitudinal dark stripes on the lower jaw, on the first two or three in- 
fralabials and the edges of the chin shields. 

Differences in proportions as expressed in number of ventrals and sub- 



228 Proceedings of the Biological Society of Washington 

caudals and relative length of tail here support the differences shown in 
the coloration (e. g., males of this population have tails considerably 
longer than those of the other two subspecies), and these data will be 
presented later in a detailed consideration of variation within the species. 

Description of Holotype: Top of head with the nine symmetrically 
arranged scutes characteristic of the genus but with modifications as fol- 
lows: a small triangular scute at each anterolateral point of the frontal; 
a narrow elliptical scale between the frontal, left supraocular, and left 
parietal; a single triangular scale occupying the notch between each 
parietal and its corresponding supraocular. Each parietal truncated by a 
transverse suture separating the posterior third. Two nasals, the posterior 
extending forward into the nostril. No loreals. Two preoculars, the lower 
one smaller, forming the upper posterior border of the pit, and having a 
suture-like groove across the posterior tip near the orbit. A very small 
postfoveal entering the orbit on each side. Two postoculars, the upper 
one small, the lower crescent-shaped and forming the lower posterior 
border of the orbit. Supralabials 8, the second extending upward to form 
the anterior border of the pit, the third largest and extending upward to 
form the lower anterior border of the orbit. A small postfoveal forms the 
lower posterior border of the pit. Infralabials 10 on the left side, 11 on 
the right. Temporals in five longitudinal rows at a point opposite the 
posterior ends of the parietals; the scales of the lowest row larger, and 
the anterior two of these without keels. The posterior tips of the first 
infralabials extend backward between the chin shields of which there 
is one pair. Four rows of median gulars, the two inner ones less than 
half the width of the outer ones; five or six irregular rows of lateral 
gulars. 

Dorsal scales each with two conspicuous apical pits, and all strongly 
keeled on body and tail. Scale rows 27—25-23-21, the order of decrease 
(Dowling recount system, 1951) as follows: 

-7(12) -6(107) -6(129) 

27 25 23 21 (14D0 

-7(12) -6(107) -5(130) 

Ventrals 141, plus a half one on the right side. Anal plate not divided. 
Both hemipenes everted. Subcaudals 50, the proximal 34 and the 37th 
single (undivided), the 35th, 36th, and the distal 13 in pairs (divided). 
There is a scar from a wound in the skin on the right side opposite ven- 
trals 41-45. Total length 795 mm; tail length 142 mm; tail/total length 
ratio .178. 

Top of head dark brown in frontal and parietal region, pale brown 
on outer margins of internasals, prefrontals, and supraoculars. Muzzle 
generally light. Sides of head, lower jaws, and throat cream to white and 
conspicuously marked with dark brown as follows: sharply defined ver- 
tical bars on the sides of the rostral and the adjacent edges of the pre- 
nasals and first supralabials; a short dark stripe from the posterior edge 
of the postnasal obliquely downward across the preoculars and postfoveal 



Two new snake subspecies 229 

to the orbit; a broad dark cheek stripe, sharply defined above and below 
with white, extending obliquely backward from the orbit and beyond 
and beneath the angle of the jaw, covering the upper edges of the supra- 
labials, the lowest row of temporals, and the lower halves of the tem- 
porals of the second row; a triangular spot at the sutures between the 
1st and 2nd, 3rd and 4th, and 4th and 5th supralabials; a dark stripe 
from the mental obliquely backward across infralabials 1 to 5 and the 
outer edges of the chin shields; and dark spots at the sutures of infrala- 
bials 5 and 6, 6 and 7, and 8 and 9. 

Dorsal coloration of body pale brown with 12 dark brown crossbands, 
the ground color becoming darker posteriorly and dark brown to black 
on the tail. The crossbands are 8 to 11 scales wide on the sides (on scale 
rows 4 or 5) and 6 to 8 scales wide middorsally; they are dark brown at 
the margins, pale brown in the central portions, which here are only 
slightly darker than the ground color, and contain one or more irregular 
dark blotches on the sides. The margins of the crossbands extend onto 
the ventrals as dark areas to correspond with a separate series of small 
blotches and thus form an irregular row of dark ventrolateral spots on 
each side. 

The tail is generally dark brown to black above and below, except the 
distal half which is cream to white below; it is crossed by approximately 
10 indistinct dark bands. The belly is cream-colored anteriorly, but this 
becomes obscured by dark pigmentation toward the tail. 

Notes on Paratypes: The seven paratypes range in size from a subadult 
male (total length 396 mm) to a moderately large adult female (1055 
mm). All show the sharply defined diagnostic pattern of the head. In 
structural characters there is only slight variation: supralabials 8-8 in 
all; infralabials 10 or 11; preoculars 2-2 with a small postfoveal entering 
the orbit; postoculars 2-2 or 3-3, in one instance 3-4. Ventrals, two 
males, 136, 141; five females, 132-138 (136); subcaudals, males, 47, 
50; females, 42-46 (44); divided subcaudals, males, 13, 16; females, 
12-24 (18). Tail/total length ratio, males, .169, .183; females, .158-.178 
(.161). 

A body pattern of broad crossbands is distinct in all, but in the 
largest of the series it becomes obscured posteriorly by darker pigmenta- 
tion which also involves the ground color. Crossbands 13 in males, 11 to 
13 in females, 4 to 8 scales wide at the midline, 7 to 11 scales wide on the 
sides; darker at the edges, with central areas little different in tone from 
the ground color and usually with dark blotches of varying size and in- 
tensity. Occasionally irregular half-bands alternate instead of meeting 
evenly at the midline. Both ground color and crossbands become pro- 
gressively darker posteriorly. In all except the subadults the crossband 
pattern of the tail is obliterated, and with increasing size (age) the 
tail becomes black above and below. When the pattern of the tail can 
be seen, there are 8 or 9 irregular crossbands and the tip is light, some- 
times with a trace of yellow. The ventral ground color is white, cream, 
or pale tan, darker in younger individuals. The ventral markings are 



230 Proceedings of the Biological Society of Washington 

blotches or spots o£ reddish brown, brownish black, or black, stronger 
toward the sides, more distinct in younger individuals, and usually 
flowing together posteriorly into a nearly continuous black area in older 
ones. 

This subspecies is named in honor of a friend and colleague, Roger 
Conant, Director of the Philadelphia Zoological Garden, whose profes- 
sional achievements and dynamic promotion of the study of herpetology 
for amateurs merit highest commendation. 

Geographic distribution: The range of A. p. conanti includes all of 
Florida and its off-shore islands and the southern part of Georgia. In- 
tergradation with A. p. leucostoma occurs in southwestern Alabama, and 
with A. p. piscivorus in southeastern Alabama and southern and south- 
eastern Georgia. 

Key to the Subspecies of the Cottonmouth 

1. Tip of the snout conspicuously marked by a pair of dark vertical 
lines at the edges of the rostral and the adjacent prenasals and first 
supralabials ( Fig. 6 ) ; markings of the head usually sharply defined, 
except in some large (old) individuals; tail relatively long, aver- 
aging 18 percent of the total length in males, 16 percent in females; 

subcaudals 45-54 (50) in males, 41-49 (45) in females 

conanti, new subspecies. 

See Geographic distribution, above. 

Tip of the snout without dark vertical lines at the edges of the 
rostral, adjacent prenasals and first supralabials; markings of the 
head usually not sharply defined except in juveniles; relative length 
of the tail averaging 16 percent in males, 15 percent in females 2 

2. Top and sides of the snout dark brown or black with no visible 
pattern; general coloration relatively dark, even in juveniles; sub- 
caudals 38-49 (44) in males, 36-48 (42) in females 

leucostoma ( Troost), 1836. 

The Mississippi Valley from southeastern Kansas, central Missouri, 
southern Illinois, western Kentucky, and western Tennessee south 
to the Gulf of Mexico; west to central Oklahoma, west central Texas, 
and along the Gulf Coast from the vicinity of Corpus Christi Bay 
east to southern Alabama. 

Top and sides of the snout generally light brown, no pattern of 
lines or stripes; general coloration relatively light; crossbands of 
body usually in strong contrast with the ground color; subcaudals 

39-51 (46) in males, 41-50 (44) in females 

piscivorus ( Lacepede ) , 1789. 

Atlantic Coastal Plain from east-central Alabama and eastern Geor- 
gia to southeastern Virginia. 

Acknowledgments: For assistance in obtaining living specimens from 
selected localities in Kansas and Florida I am indebted to Dr. Henry S. 
Fitch, Dr. William J. Riemer, Dr. Walter Auffenberg, and WilHam G. 






Two neiv snake subspecies 231 

Weaver, Jr. During the many years of our collaboration and frequent 
correspondence Roger Conant has provided stimulating discussion and 
encouragement; his long-standing interest in this project is sincerely 
appreciated. Dr. Hobart M. Smith has responded most helpfully to my 
requests for advice on matters of procedure. This paper includes results 
of part of a study supported by the National Science Foundation ( G2896, 
G8702, andG19400). 

Names of Museums Abbreviated in Text 

AMNH — American Museum of Natural History, New York 

CA — Chicago Academy of Sciences 

FMNH — Field Museum of Natural History, Chicago 

KU — University of Kansas, Museum of Natural History, Lawrence 

UCM — University of Colorado Museum, Boulder 

UMMZ — University of Michigan Museum of Zoology, Ann Arbor 

USNM — United States National Museum, Washington, D. C. 

Literature Cited 

Beauvois, Palisot de, a. M. F. J. 1799. Memoir on Amphibia. Ser- 
pents. Trans. Amer. Philos. Soc, 4: 362-381, 1 pi. 

DowLiNG, Herndon G. 1951. A proposed method of expressing scale 
reductions in snakes. Copeia, 1951, 2: 131-134. 

Gloyd, Howard K. and Roger Conant. 1934. The broad-banded 
copperhead: a new subspecies of Agkistrodon mokasen. Occ. 
Papers Mus. Zool. Univ. Michigan, 283: 1-5, pi. 1. 

. 1943. A synopsis of the American forms of Agkistrodon 

(copperheads and moccasins). Chicago Acad. Sci. Bull., 7 
(2): 147-170, fig. 1-16, 2 maps. 

Klauber, Laurence M. 1948. Some missapplications of the Linnaean 
names applied to American snakes. Copeia, 1948, 1: 1-14. 

Lacepede, Compte de. 1789. Histoire naturelle des serpens. Paris: 
Hotel de Thou, vol. 2, 527 p. 

Linnaeus, Carolus. 1766. Systema naturae . . . ed. 12. Holmiae: 
Laurentii Salvii, vol. 1, 532 p. 

Smith, Hobart M. and Howard K. Gloyd. 1964. Nomenclatural 
notes on the snake names Scytale, Boa scytale, and Agkistro- 
don mokasen. Herpetologica, 19 (4): 280-282. 

Troost, G. 1836. On a new genus of serpents, and two new species of 
the genus Heterodon, inhabiting Tennessee. Ann. Lye. Nat. 
Hist. New York, 3: 174-190, pi. 5. 



232 Proceedings of the Biological Society of Washington 



7i 



QH 
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B4X 
IMH 


;33-242 


PROCEEDINGS 
OF THE 


3 October 1969 


BIOLOGICAL 


SOCIETY OF WASHINGTON 



THE RED-THROATED ANT-TANAGER ( HABIA^^go/V/^Ns 
FUSCICAUDA) IN PANAMA AND COLOMB^J^^ ^^ 

By Kenneth C. Parkes V ^^^ ?' '™^ 

Carnegie Museum, Pittsburgh, Pennsylvania\^^(^iQnf.o\^:» 

The Red-throated Ant-tanager ( Habia fuscicauda ) is princi- 
pally a species of Middle America (see map in Willis, 1960: 
151). It was formerly considered to be conspecific with the 
isolated Sooty Ant-tanager (H. gutturalis) of the middle Mag- 
dalena and other valleys of the Colombian Andean region 
(Meyer de Schauensee, 1966: 484-485), and much of the 
literature of the Red-throated Ant-tanager appears under the 
specific name Habia gutturalis. Both morphological and be- 
havioral evidence {fide E. O. Willis) indicate that gutturalis 
and fuscicauda are best kept as separate species. 

At the southern end of its range H. fuscicauda appears to 
have a rather interrupted distribution, in eastern Panama and 
Colombia. Meyer de Schauensee (1966: 485) gives the South 
American range as follows: "COLOMBIA, Caribbean coastal 
region west of the rio Magdalena from Atlantico to Cordoba 
(upper Sinu valley), possibly Santa Marta but no authentic 
records." Carriker (1955: 60) doubted the Atlantico "sight" 
record (Santa Cruz, Los Pendales) that had earlier been ac- 
cepted by Meyer de Schauensee (1951: 1054), although 
Carriker conceded that the species might occur north to the 
hills of central Atlantico. Hellmayr (1936: 314) examined a 
specimen alleged to have been taken at Barranquilla, Atlantico, 
but Meyer de Schauensee (1951: 1054) placed this locality in 
quotation marks, presumably an indication that he believed the 
"Barranquilla" specimen to have been a native trade skin of 
uncertain origin. 

19— Proc. Biol. Soc. Wash., Vol. 82, 1969 (233) 



234 Proceedings of the Biological Society of Washington 

The name Phoenicothraupis erythrolaema Sclater, 1862, was 
based on a manuscript name of Bonaparte, attached to speci- 
mens in the Verreaux collection attributed to "S. Martha, New 
Grenada" (= Santa Marta, Colombia). Sclater compared his 
erythrolaema only to ''Phoenicothraupis rubicoides," which is 
a Mexican subspecies of another species, Habia rubica. Sclater's 
name was placed in the synonymy of Phoenicothraupis fusci- 
cauda Cabanis (Costa Rica) by Lawrence (1863: 9), where it 
remained until revived by Bangs (1900: 30). In the latter 
paper Bangs attributed to erythrolaema a range "extending 
from Santa Marta to Panama," but later Thayer and Bangs 
(1906: 222) expressed some doubt as to the correctness of the 
alleged provenience of the type, as no ant-tanagers had been 
collected subsequently in the Santa Marta region. Todd and 
Carriker (1922: 483) agreed, and considered the Santa Marta 
locality to be almost certainly erroneous. They did not, how- 
ever, venture to conjecture about the actual origin of the type 
of eiythrolaema, mentioniug merely that the name had been 
applied to the Panama form by Bangs. 

As a brief digression, it should be mentioned that since 1922 
the generic name Habia Blyth, 1840, has been used for the ant- 
tanagers in place of Phoenicothraupis Cabanis, 1851. The lat- 
ter generic name (and, in fact, its root Thraupis) has been 
variously treated in the literature as being of masculine or 
feminine gender. To avoid confusion, all names in the present 
paper, except when quoted directly, will be treated as femi- 
nine, in agreement with the current generic name Habia. 

In January 1916 Carriker collected for Carnegie Museum a 
series of ant-tanagers near Turbaco, Bolivar, Colombia, just 
inland from Cartagena. Tood (1917), in a brief "preliminary 
diagnosis," described the Turbaco birds as a new species, 
Phoenicothraupis rubiginosus. He compared the male with 
''Phoenicothraupis salvini salvini" ( = Habia fuscicauda salvini, 
type locality Vera Paz, Guatemala ) , and the female with Costa 
Rican fuscicauda. He did not mention erythrolaema. Todd's 
unpublished notes, filed at Carnegie Museum, go into more de- 
tail than did his formal published diagnosis of "rubiginosus." 
He dismisses the name erythrolaema from consideration for his 



J 



The Red-throated Ant-tanager 235 

new Colombian species, partly on the basis that the type locality 
Santa Marta was probably erroneous, and partly because sev- 
eral authors had considered males of eiythrolaema inseparable 
from fuscicauda, whereas in Todd's opinion males of "rubigi- 
nosus" were barely separable from salvini. Todd did not list his 
comparative material of salvini, but it was presimiably the 
series from British Honduras then available in Carnegie Mu- 
seum, which is indeed best referred to salvini, contra Russell 
(1964: 173). Later Todd added to his notes the following 
statement: "Panama skins (erythrolaema) prove upon com- 
parison to be so exactly intermediate in all respects between 
fuscicauda and rubiginosa that the only course open is to make 
all three forms conspecific." These observations are undated, 
and were probably made independently of those of Dwight 
and Griscom (1924: 5). In any case, the latter authors were 
the first to point out in print that three forms were possibly in- 
volved: fuscicauda of Costa Rica; erythrolaema from "some- 
where on the north coast of Colombia," with rubiginosa Todd 
as a synonym; and an unnamed Panamanian race of fus- 
cicauda, to which the name "erythrolaema" had been errone- 
ously applied by Thayer and Bangs. In the principal standard 
reference work on New World birds, Hellmayr (1936: 314) 
briefly reviewed this history, and synonymized rubiginosa Todd 
with erythrolaema, on the basis that he could not separate an 
adult male from Baranquilla from a series from along the Pan- 
ama Railroad (but see above on the status of this "Barran- 
quilla" specimen). Hellmayr, in a lengthy footnote, para- 
phrased a letter from Todd concerning the latter's opinion of 
the validity of rubiginosa after having examined specimens at 
the British Museum. Hellmayr s wording is ambiguous, and it 
is not always clear whether Hellmayr or Todd is responsible 
for a given statement. I therefore consulted the carbon of 
Todd's original letter to Hellmayr, dated 1 August 1934. Todd's 
findings were (1) as he had already noted on his file card, 
his rubiginosa was not a good species but was conspecific with 
fuscicauda of Costa Rica; (2) Panama specimens were "clearly 
intermediate" between those of Costa Rica and those of Colom- 
bia, for which he advocated use of his name rubiginosa; (3) 



illHTHSUNlA^ JSPJ3 1969 



236 Proceedings of the Biological Society of Washington 

the type of erythrolaema. was "rather faded" but appeared to 
belong to the Panama race; (4) the name erythrolaema was 
therefore available for the Panama birds, but Todd's inclina- 
tion was to recognize only the extreme forms fuscicauda and 
rubiginosa, suppressing the name erythrolaema entirely. Hel- 
Imayr obviously disagreed with these findings, preferring to 
unite Panamanian and Colombian birds under the older name 
erythrolaema. 

No further developments in the nomenclatorial history of 
these ant-tanagers took place until 1955, when Carriker desig- 
nated the type locality of "erythrolaemus" [sic = erythrolaema] 
as Turbaco, Colombia ( where he had collected ant-tanagers in 
1916), stating that the type specimen probably came from near 
Cartagena. Carriker inadvertently gave the original supposed 
origin of the type as "Magdalena"; as stated above, the type lo- 
cality was given as "S. Martha," or Santa Marta. This action of 
Carriker s would make ruhiginosa Todd an objective synonym 
of erythrolaema Sclater, unless it could be demonstrated that 
the type of erythrolaema probably did not come from the vicin- 
ity of Cartagena ( International Code of Zoological Nomencla- 
ture, 1964, Recommendation 72E). Carriker did not examine 
the type of erythrolaema, and accepted without question its 
identity with the birds of Turbaco. 

The suggestion of D wight and Griscom (1924: 5) that the 
Panamanian population of fuscicauda might need a name was 
never pursued, all subsequent writers having accepted Hell- 
mayr's usage. As pointed out by Carriker (1955), the only 
modem specimens from Caribbean Colombia, and the only 
ones with accurate locality data, are those he himself collected 
in 1916 and 1942. The former series, as stated above, is in 
Carnegie Museum, the latter in the Smithsonian Institution. 
Far more material is now available than was true when Hell- 
mayr, Todd and other previously mentioned writers studied 
these ant-tanagers. Central to the problem remained the status 
of the name erythrolaema. Through the courtesy of Dr. David 
Snow I was able to borrow Sclater s type specimen, in order to 
try to settle the matter once and for all. 

Upon comparison of the type of erythrolaema ( BM 1885-6- 



The Red-throated Ant-tanager 237 

12-681) with Panamanian birds and with the type series of 
rubiginosa, I disagree with Todd on two counts. This speci- 
men, although now well over a century old, does not appear to 
me to be "rather faded" — no more so than the now 53-year-old 
specimens of rubiginosa, or a 69-year-old Panamanian specimen 
at hand. And I do not consider it to be "identical" ( Hellmayr's 
word ) with the Panamanian birds. 

The type of erythrolaema is an almost perfect match for CM 
52413, the only specimen in the type series of rubiginosa that 
was not taken at Turbaco. It was collected by Carriker at 
Puerto Zapote, Bolivar, which ( according to Meyer de Schau- 
ensee, 1948: 324) is the same as Cispata, on the east side of 
Cispata Bay, opposite the mouth of the Rio Sinu, or about 125 
kilometers farther southwest along the coast ( toward Panama ) 
than Turbaco. Both the type and the Cispata bird are some- 
what intermediate between the Turbaco series and the Pana- 
manian form. In view of the extensive deforestation that has 
taken place all along this part of Caribbean Colombia, it is 
impossible to say whether this species may not once have had 
an essentially uninterrupted range from eastern Panama to the 
vicinity at least of Cartagena; the intermediacy of the Cispata 
bird at least suggests that there has been gene flow. Since the 
type of erythrolaema does not match Turbaco birds, I would re- 
ject Carriker s designation of Turbaco as the type locality, and 
substitute "vicinity of the mouth of the Rio Sinu"; the specimen 
was probably obtained from a professional collector, or native 
plumage hunters, for the Verreaux collection. Specimens of this 
era labelled "Santa Marta" (or one of the variant spelHngs) 
could have come from any of several different parts of Colom- 
bia (Todd and Carriker, 1922: 21-22). The type locality, 
therefore, may best be fixed on the basis of the characters of 
the type specimen, which, as stated above, match the bird 
from the mouth of the Rio Sinu. 

Although erythrolaema is therefore not absolutely identical 
with Todd's rubiginosa, I would recommend using the name 
erythrolaema for all of the Caribbean Colombian Habia fusci- 
cauda, considering rubiginosa as a synonym. Contrary to Todd, 
I find the type of erythrolaema nearer the Turbaco series than 



238 Proceedings of the Biological Society of Washington 

to those of Panama in color. In the Hght of the extensive de- 
forestation, it will be interesting to see whether any remnant 
populations of this species are rediscovered in what is now the 
apparent distributional gap between eastern San Bias, Panama, 
and the Rio Sinu in Colombia. One would predict that such 
birds might well represent additional intermediates, stages 
along a former cline between the extreme ( "rubiginosa" ) type 
of the Turbaco region and the "typical" Panamanian birds of 
central Panama. 

The Panamanian birds remain to be discussed. In spite of the 
fact that they have hitherto been included nomenclatorially 
with the Colombia birds, those of Panama are (as predicted 
by Dwight and Griscom, 1924) a perfectly recognizable sub- 
species, rather more like Costa Rican fuscicauda than like the 
Turbaco birds. The Panama birds may be known as: 

Habia fuscicauda willisi new subspecies 

Holotype: United States National Museum no. 445515, adult $ , from 
Boca del Rfo Indio, Colon, Panama, collected by Alexander Wetmore and 
W. M. Perrygo on 15 February 1962 (original number 16757). 

Characters: Definitively plumaged males differ from erythrolaema as 
follows: throat deeper and richer red; throat/breast color transition more 
abrupt ( whereas in erythrolaema there is a fairly abrupt breast/ abdomen 
transition); breast and abdomen much darker and grayer; flanks heavily 
washed with gray; midventral region and under tail coverts dark red 
with either a scarlet or bluish tinge, not rose or orange; black spot of 
malar region averaging more extensive, occupying much of subocular 
region; ear coverts darker; crest feathers somewhat more elongated; crown 
patch larger, darker and more intensely red; dorsum darker, more pur- 
plish red; wings and tail much darker, almost black in some individuals. 
Females differ from erythrolaema as follows: throat patch deeper, more 
intense yellow, abruptly contrasting with breast rather than blending 
with breast color; posterior underparts very much darker — dark, dull 
olive brown with a yellowish wash medially, rather than grayish yellow, 
somewhat browner on the flanks as in erythrolaema; under tail coverts 
darker and browner; ear coverts dark olive brown rather than light yel- 
lowish brown; back, wings, and tail all darker. 

Definitively plumaged males differ from fuscicauda as follows: throat 
more brilliantly red; posterior underparts, on the average, less shaded 
with gray, the red tending toward pinkish or bluish rather than toward 
brownish or orange; dorsum averaging lighter and brighter, of either a 
purer or bluer, less brick red; crest purer, less orange red; tipping of 
crest feathers in fresh plumage browner, less black. Females differ from 



The Red-throated Ant-tanager 239 

fuscicauda as follows: throat deeper, more brilliant yellow; little or no 
brownish band across breast; posterior underparts paler, less washed with 
dark brown; abdomen with much more yellow wash; dorsum less brown- 
ish, more greenish; crown more greenish yellow, less ochraceous yellow. 

Range: central Panama, chiefly on the Caribbean slope. Specimens 
were examined from the provinces of Colon, northeastemmost Code, 
Panama, and westernmost San Bias (Mandinga), as well as from the 
Canal Zone. Western Bocas del Toro province (Almirante and vicinity) 
is inhabited by typical fuscicauda. A series of 9 specimens in the AMNH 
collection from Santa Fe, northern Veraguas, is highly variable, and 
represents intergradation between fuscicauda and willisi. Santa Fe is just 
about halfway between Almirante and the type locality of willisi. Eugene 
Eisenmann has kindly suppHed the following information: "In Panama 
this species is primarily a bird of the humid Caribbean slope, though it 
has crossed over to the Pacific slope at low divides in Veraguas, as at 
Santa Fe, and eastward in Panama province and the Canal Zone, where 
sufficiently humid conditions and shady woodland exist." The apparent 
gap in the range of the species between northwestern Bocas del Toro and 
central Colon (except for the one series from Santa Fe) is interpreted 
by Eisenmann as due partly to inadequate collecting and partly to un- 
suitably mature forest with too little undergrowth. As in the case of the 
bird from the Rio Simi in Colombia, the intermediacy of the Santa Fe 
specimens suggests populations in sufficient contact for gene exchange. 

From the Pacific slope there are specimens from the Canal Zone and 
from adjacent Panama province east to the Rio Pacora, about 35 miles 
east of the city of Panama (USNM — Wetmore). West of the Canal Zone 
and east of Santa Fe I have seen only a single first-year male in Carnegie 
Museum collected by A. C. Twomey at Capira, Panama province. Both 
Willis and Eisenmann have observed this species (with army ants) on 
Cerro Campana, a few miles beyond Capira, which Eisenmann's notes 
characterize as "a low mountain right on the divide . . . extremely humid 
at the upper levels." 

Hellmayr (1936: 314) listed a supposed record of "Phaenicothraupis 
fuscicauda" from the Rio Lara, Darien, based on a specimen collected by 
Festa in 1895 and pubHshed by Salvador! and Festa (Bol. Mus. Zool. 
Anat. Comp. R. Univ. Torino, 14, no. 339, 1899: 4 — original publication 
not seen). Dr. Wetmore (letter of 16 January 1969) beheves this record 
to be erroneous and probably based on a specimen of Habia ruhica. As- 
suming Dr. Wetmore to be correct in this assumption, there are no speci- 
men records of Habia fuscicauda east of the Rio Pacora and Rio Mand- 
inga in Panama or west of the Rio Simi in Colombia. One sight record 
from the intervening area is believed by Eisenmann to be worthy of ser- 
ious consideration. According to information suppHed to me by Eisen- 
mann, Dennis R. Sheets reported "several pairs heard, one seen" low 
down in overgrown thickets above Puerto Obaldia, on the trail to La 
Bonga, 16-18 June 1965. Puerto Obaldia is in easternmost San 



240 Proceedings of the Biological Society of Washington 

Bias, virtually on the Colombian border, and thus just about halfway be- 
tween the extreme known specimen localities given above. Eisenmann 
informs me that Sheets was well acquainted with this ant-tanager 
in the Canal Zone, and in view of the highly distinctive nature 
of its vocalizations it is unlikely that another species was mistaken by 
Sheets for H. fuscicauda. The superficially similar Red-crowned Ant- 
tanager (Habia rubica), which is quite different in voice from H. fusci- 
cauda, is not known from the Caribbean slope of Panama (fide Eisen- 
mann ) . 

Etymology: it is most appropriate that this subspecies of ant-tanager 
should be named for Dr. Edwin O. Willis, who has contributed so much 
to our knowledge of the biology of the tanagers of the genus Habia. 

Specimens examined: This study was based on the combined series of 
the pertinent forms belonging to Carnegie Museum, the United States 
National Museum, and the American Museum of Natural History, plus a 
series from the Museum of Comparative Zoology, and the type of ery- 
throlaema. The total number examined is as follows: H. /. fuscicauda, 42 
from Costa Rica and 11 from Panama; H. f. fuscicauda X willisi, 9 from 
Santa Fe, Veraguas, Panama; H. f. willisi, 121 from Panama; H. /. 
erythrolaema, 21 from Colombia. 

Thayer and Bangs (1906: 222) claimed that specimens from the sa- 
vannas near the city of Panama, on the Pacific slope, were the same as 
erythrolaema, and more pallid than those from Loma del Leon (in the 
area now occupied by the Canal Zone). The latter they ascribed to 
"true P. fuscicauda.'* I have examined the two series involved, and find 
that the savanna series does, on the whole, average slightly paler. This 
may well be attributable, at least in part, to the fact that they are May 
specimens, and thus more worn than the March birds from Loma del 
Leon. The minor differences between the two series are bridged by in- 
dividual variation, which tends to be strong in this species, and all may 
be called willisi. Loma del Leon birds are not fuscicauda nor are the 
savanna birds erythrolaema. 

Acknowledgments: I am indebted to the authorities of the Smithsonian 
Institution and the American Museum of Natural History for access to 
their collections. Dr. David Snow of the British Museum (Natural His- 
tory) was good enough to arrange to have the type of Phoenicothraupis 
erythrolaema sent on loan. Dr. Raymond A. Paynter, Jr., lent the Panama 
specimens from the Museum of Comparative Zoology that had been dis- 
cussed by Thayer and Bangs. Dr. Alexander Wetmore allowed me to 
use collections of Panamanian and Colombian birds set aside for his per- 
sonal research, and provided additional useful information about the 
distribution of Habia fuscicauda in those countries. Finally, Mr. Eugene 
Eisenmann kindly extracted from his notes on Panama birds everything 
pertaining to the distribution and habitats of ant-tanagers in that country, 
and made this information freely available to me. 



The Red-throated Ant-tanager 241 

Literature Cited 

Bangs, O. 1900. List of birds collected by W. W. Brown, Jr., at Loma 
del Leon, Panama. Proc. New England Zool. Club, 2: 13^34. 

Carriker, M. a., Jr. 1955. Notes on the occurrence and distribution 
of certain species of Colombian birds. Novedades Colom- 
bianas, 1: 48-64. 

DwiGHT, J., and Griscom, L. 1924. Descriptions of new birds from Costa 
Rica. Am. Mus. Novitates, no. 142: 5 pp. 

Hellmayr, C. E. 1936. Catalogue of birds of the Americas and adjac- 
ent islands. Part 9. Field Mus. Nat. Hist. Zool. Ser., 13, 
part 9: V + 458 pp. 

Lawrence, G. N. 1863. Catalogue of a collection of birds made in New 
Granada, by James McLeaiman, Esq., of New York, with 
notes and descriptions of new species. Part IV. Aim. Lye. 
Nat. Hist. New York, 8: 1-13. 

Meyer de Schauensee, R. 1948. The birds of the Republic of Colombia. 
Part I. Caldasia, 5: 251-380. 

. 1951. Ibid., Part 4. Caldasia, 5: 873-1112. 

. 1966. The species of birds of South America and their dis- 
tribution. Acad. Nat. Sci. Philadelphia: xvii + 577 pp. 

Russell, S. M. 1964. A distributional study of the birds of British Hon- 
duras. Omith. Monog. no. 1, Am. Ornith. Union: 195 pp. 

Sclater, p. L. 1862. Catalogue of a collection of American birds belong- 
ing to Philip Lutley Sclater, M.A., Ph.D., F.R.S. London, N. 
Trubner & Co.: xiv + 368 pp. 

Thayer, J. E., and Bangs, O. 1906. Vertebrata from the Savanna of 
Panama. III. Aves. Bull. Mus. Comp. Zool, 46: 213-224. 

Todd, W. E. C. 1917. Preliminary diagnoses of apparently new birds 
from Colombia and Bolivia. Proc. Biol. Soc. Washington, 30: 
3-6. 

, and Carriker, M. A., Jr. 1922. The birds of the Santa 

Marta region of Colombia: a study in altitudinal distribution. 
Ann. Carnegie Mus., 14: 3^582. 

Willis, E. O. 1960. A study of the foraging behavior of two species of 
ant-tanagers. Auk, 77: 150-170. 



242 Proceedings of the Biological Society of Washington 



73 

QH 

1 

B4X 3-280 3 October 1969 

HH 

PROCEEDINGS 

OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 

TWO NEW GENERA AND TWENTY-TWO NEW^-^^.^^^ 
SPECIES OF CRABS FROM JAPAN /^^ ^^N^^ 

By Tune Sakai ( OCT 20 1969 i^ 

Biological Department, Yokohama Nationar< 

University, Yokohama, Japan ^ ^ i ARkR ^ , | 3^ 

During the past two or three years, a considerable number 
of crab specimens from the central and southern districts of 
Japan have been sent to the author for identification or as 
gifts. Collections were made either along the shore, from the 
refuse of lobster nets in shallow water or from fishing trawlers 
working on the continental shelf. The main sources of materials 
were: a collection from S. Yamamoto and pupils (Nagashima 
High School) along the coast of Kii Nagashima, Mie Prefecture, 
obtained from lobster nets; the collection of the late Mitsuno- 
suke Ozaki, which includes more than 300 species collected by 
him along the coast of Kii Minabe, Wakayama Prefecture, dur- 
ing his 20 years of work; Katsushi Sakai's collection from off 
Mimase, Tosa Bay, which includes many previously unrecorded 
species from Japan, mostly obtained by trawling; Miyoshi 
Matsuo's collection from the mud flat of Ariake Bay, Nagasaki 
Prefecture; and a collection from Katsushi Sakai and Hirosuke 
Yamada from the coral reef of Yoron Island, the southernmost 
island of the Amami Group. 

Besides these, some new species were donated by Dr. T. 
Odawara and Mr. H. Hayashi of the Odawara Carcinological 
Museum, Tokyo, and some species were obtained by the author 
himself during a collecting trip to Mikawa Province, Kii Pe- 
ninsula, Tosa Bay and Ariake Bay. 

The present paper describes the following 22 new species 
belonging to 11 families and 21 genera, 2 of which are new: 

Fam. DROMIIDAE 

Cryptodromia cristatipes new species 

20— Proc. Biol. Soc. Wash., Vol. 82, 1969 (243) 



244 Proceedings of the Biological Society of Washington 



Fam. LEUCOSIIDAE 

Arcania sagamiensis new species 

Leucosia mimasensis new species 
Fam. HYMENOSOMIDAE 

Neorhynchoplax ariakensis new species 
Fam. MAJIDAE 

Epialtus orientalis new species 

Sphenocarcinus hidens new species 

Chlorinoides tosaensis new species 

Leptomithrax kiiensis new species 

Maja nagashimaensis new species 
Fam. PARTHENOPIDAE 

Parthenope (Pseudolambrus) ozakii new species 
Fam. CANCRIDAE 

Cancer nadaensis new species 
Fam. PORTUNIDAE 

Benthochascon elongatum new species 

Thalamita yoronensis new species 
Fam. XANTHIDAE 

Neoliomera acutidens new species 

Neoliomera richteroides new species 

Planopilumnus minabensis new species 

Heteropilumnus mikaivaensis new species 
Fam. GONEPLACIDAE 

Carcinoplax tomentosa new species 

Psopheticoides sanguineus new genus and species 
Fam. PINNOTHERIDAE 

Orthotheres turhoe new genus and species 

Sakaina incisa new species 
Fam. OCYPODIDAE 

Macrophthalmus (Macrophthalmus) ceratophorus 
new species 

In the course of the study, the author visited numerous mu- 
seums, universities and institutions in order to compare the 
specimens with types of related species. For the privilege of 
examining material and referring to literature, as well as for 
kind advice, the author wishes to extend his thanks to: 

Dr. R. Serene at the National Museum, Singapore; Drs. J. 



New Japanese crabs 245 

Forest and D. Guinot of the Museum d'Histoire Naturelle, Paris; 
Dr. L. B. Holthuis of the Rijksmuseum van NatuurHjke His- 
toric, Leiden; Dr. T. Wolf of the Copenhagen Museum, Den- 
mark; Dr. I. Gordon of the British Museum (Natural History), 
London; Dr. W. L. Schmitt, Dr. F. A. Chace, Jr., Dr. H. B. 
Roberts of the Smithsonian Institution, Washington D.C.; Dr. 
F. M. Bayer and Dr. A. J. Provenzano of the Institute of Marine 
Sciences, Miami; Dr. J. S. Garth of the Allan Hancock Founda- 
tion, Los Angeles; and Dr. D. M. Devaney of the Bernice P. 
Bishop Museum, Honolulu, Hawaii. 

The author also wishes to acknowledge especially the various 
collectors of new species for their kindness in donating the 
specimens. 

The holotypes of the new species described in this paper 
are deposited in the Smithsonian Institution ( SI ) , Washington, 
D.C., U.S.A. 

Family Dromiidae Alcock 

Genus Cryptodromia Stimpson 

Cryptodromia cristatipes new species 

PI. I, fig. 1 

Five ^ ^ , 3 $ $ , one male ( SI 125866 ) of which is designated as 
the holotype. Tosa Bay, depth 150 m. Coll. by K. Sakai. 

Two $ $. Off Mikawa Bay, Aichi Prefecture, depth 100 m. Coll. 
by H. Hayashi of the Odawara Carcinological Museum, Tokyo. 

One ^ , 1 $ . Kii Nagashima. Coll. by S. Tanaka of the Board of 
Education, Mie Prefecture. 

One S,l 2 . Kii Minabe. Coll. by the late M. Ozaki. 

Diagnosis: The carapace is sub-quadrilateral, a little broader than long. 
The dorsal surface is moderately convex, the regions being rather ill- 
defined and thickly covered with fine studded granules. The front pro- 
trudes strongly forward beyond the outline of the carapace; its free 
margin is markedly cristate and the median portion strikingly depressed, 
forming a tiny low triangular tooth. The lateral frontal margin is com- 
pletely continuous with the outer orbital angle. The lower orbital edge is 
granulated, and its inner angle formed into a strong process. 

The lateral borders of the carapace are subparallel. Anteriorly they are 
continuous with the antero-extemal angle of the buccal cavity, not with 
the outer orbital angle. Their free edge is cut into three or four laciniated 
teeth in front of the branchial groove, and also into three or four small 
ones behind the same groove. The posterolateral borders are shorter than 
the anterolateral, sHghtly concave and markedly convergent backwards. 
The posterior border is very narrow and straight. 



OCT 3 Wi 



246 Proceedings of the Biological Society of Washington 




New Japunese crabs 247 

The chelipeds are robust and much longer than any of the ambulatory 
legs, being more than twice as long as the carapace. The merus is rela- 
tively short and its outer distal surface sparingly granular. The carpus is 
robust and covered with granules. The upper distal border of the carpus 
is armed with two large processes, one at the junction with the propodus 
and the other at the antero-external angle. The propodus is also gran- 
ulated on the upper and outer surfaces, but not markedly dentate at the 
junction with the movable finger. 

The carpus of the first and second ambulatory legs is sulcate on the 
upper surface, the anterior and posterior margins being granulated. The 
third and fourth ambulatory legs are markedly reduced in size, and are 
completely dorsal in position. The last leg has a terminal pincer. 

There is no epipodite on the basis of the chelipeds. The male ab- 
domen has a platelet ( vestigial uropod ) on either side between the penul- 
timate and terminal segments. 

Measurements: Type male, length of carapace 19 mm, width of same 
22 mm. 

Relationship: The new species is closely related to Cryptodromia 
ornata Rathbun from Saya de Malha. The dorsal surface of the carapace, 
however, is less demarcated and the frontal lobe is, unlike that species, 
more strikingly cristate and its median portion only slightly dentate; in 
C. ornata the front is divided into three teeth. The type of the new spe- 
cies was compared with that of C. ornata courtesy of the Smithsonian In- 
stitution. 

Family Leucosiidae Dana 
Genus Arcania Leach 
Arcania sagamiensis new species 
Text-fig. 1 a, a' 
One $, holotype (SI 125881), 1 2, allotype. Obtained on the 
beach of Manazuru, Sagami Bay. 
Diagnosis: The carapace is subcircular in outline, a little longer than 
broad. There are ten prominent tubercles around the margin of the 
carapace, similar to the common Japanese species — A. undecimspinosa de 
Haan. The dorsal surface is thickly studded with granules and provided 

<r 

Explanation of Plate I 

Fig. 1. Cryptodromia cristatipes new species. Male holotype, dorsal 
view. X 1.4. 

Fig. 2. Maja nagashimaensis new species. Carapace of male holotype, 
dorsal view. X 1-2. 

Fig. 3. Chlorinoides tosaensis new species. Left — Male holotype, dorsal 
view. X 2. Right — Female allotype, dorsal view. X 1.4. 

Fig. 4. Leptomithrax kiiensis new species. Male holotype, dorsal view. 
X %. 



248 Proceedings of the Biological Society of Washington 





Text-fig. 1. a. Arcania sagamiensis new species. Dorsal view of male 
holotype. a'. Anterior male pleopod of same. b. Leucosia mimasensis new 
species. Dorsal view of male holotype. b'. Anterior pleopod of same ( a X 
2.5, a' X 14; b X 1.4; b' X 14). 



with fifteen tubercles, of which three are in the median line, the last one 
being on the intestinal region near the posterior border and the others dis- 
posed symmetrically on the hepatic and branchial regions. 

The front projects strikingly anteriorly and is bilobed. Each lobe ends 
in a process divided by a deep V-shaped median sinus. The outer orbital 
tooth is moderately strong. On the subhepatic region, there is a high 
tubercle, which can be seen from above just below the first marginal 
tubercle. 

The chelipeds are slender; the merus is covered with longitudinal rows 
of granules; the fingers are a little longer than the propodus and are 
finely denticulated along the prehensile edge. 

The terminal segment of the male abdomen is extremely long and nar- 
row. 

Habitat: Most of the species of Arcania are found on the bottom, 30 
to 50 meters below the surface, while this new species was obtained from 
a rocky shore in the inter-tidal zone. 



New Japanese crabs 249 

Relationship: The most closely related species is A. gracilipes Bell 
from the waters of India and Australia, but the new species has the front 
protruding more strongly anteriorly and the tubercles on the dorsal sur- 
face of the carapace are more numerous. In A. gracilipes, the cardiac 
region has two tubercles on median line, while the new species has only 
one. The type of the new species was compared with that of A. gracilipes 
by courtesy of the British Museum. 

Measurements: Length of carapace in the median line, 10 mm, width 
8.5 mm. 

Genus Leucosia Fabricius 

Leucosia mimasensis new species 

Text-fig. 1 b, b' 

One $, holotype (SI 125868), 1 2, allotype. Off Mimase, Tosa 
Bay, depth 120 m. Coll. by K. Sakai. 

Diagnosis: Related to L. obtusifrons de Haan, but this new species may 
be easily distinguished from the former by its smaller size, the peculiarity 
of the colour markings of the carapace, and the different form of the 
anterior pleopod of the male. The outline of the carapace is typically 
circular and the dorsal surface evenly convex. On either side of the gas- 
tric portion, there is a pair of dark purplish markings of a rounded trilob- 
ular shape, in each of which there are three spots of light yellow, ar- 
ranged fore and aft. The anterior spot is very small, the other two large 
and rimmed with much darker colouration. 

The front is short, its anterior edge cut obtusely. The anterolateral 
margins are rounded, obscurely lined with depressed granules near the 
junction of the antero- and postero-lateral borders. The thoracic sinus 
is very simple and circular, rimmed with about six beaded tubercles, its 
base pubescent, not tuberculated. 

The anterior male pleopod is thickened near the distal portion and 
obtusely pointed, curved inwards and furnished with a few hairs. 

Measurements: Length of carapace 17.9 mm, width of same 16.9 mm. 

Family Hymenosomidae Stimpson 
Genus Neorhynchoplax Sakai 

Most of the species of Neorhynchoplax are restricted to India and its 
adjacent regions while the species of Rhynchoplax are found in Japan and 
China with one exception — R. coralicola Rathbun, which is common to 
Japan and Thailand. Recently Mr. Miyoshi Matsuo of the Nagasaki 
Middle School obtained a very small crab living as a commensal with the 
common Holothurian, Protankyra hidentata (Woodward and Barrett), 
which inhabits the mud flat of Shimabara, Ariake Bay, Nagasaki Pre- 
fecture. This is apparently a new species of Neorhynchoplax and thus 
constitutes the first record of the genus in Japan. 



250 Proceedings of the Biological Society of Washington 




Tkxt-fig. 2. Neorhynchoplax ariakensis new species. Male holotype in 
dorsal view. ( X 9 ) 



Neorhynchoplax ariakensis new species 
Text-fig. 2; 3 a-d 

Two ^ ^, 1 2, one male (SI 125880) of which is designated as 
the holotype. Mud flat of Shimabara Peninsula, Nagasaki Pre- 
fecture. Coll. by M. Matsuo. 

Diagnosis: In its general appearance and especially in the rostral teeth, 
this new species is closely allied to Neorhynchoplax demeloi (Kemp) 
from the Mandavi River in Portuguese India. The entire body is covered 
with a soft tomentum. The carapace is rounded, slightly convex and a 
little broader than long. 

The regions of the dorsal surface are well-defined by clean-cut grooves 
as noted in the other members of the genus; the gastric region is the 
largest and is somewhat hexagonal in outline, but its anterior boundary 
is poorly defined. There are three median transverse tubercles on this 
region. The cardiac region is pentagonal with its apex directed back- 
wards, whence a short straight groove bisects the intestinal region. In 
front of the hepatic region are four clearly defined tiny facets and a very 
small orbital region. Two of the facets are located in front of the rostral 
lobes and two are on the subhepatic region. 

The true rostrum is a broad, low triangle projecting forwards. Behind 
the rostrum there are two low, obtuse lobes arranged side by side near 
the anterior boundary of the marginal groove. There are two low and 
obscure lobules on the antero-lateral borders. 

The external maxilliped has the ischium longer than the merus; the 
former is broadened distally and the latter rounded, leaving a wide hiatus 
between the opposed pair of appendages. 



New Japanese crabs 



251 





Text-fig. 3. Nearhynchoplax ariakensis new species, a. Anterior portion 
of carapace, b. Male abdomen, c. Anterior pleopod of male, d External 
maxilliped. (a X 18; b X 21; c X 60; d X 35) 



The male abdomen is sub-triangular and composed of four discrete 
segments, as represented in text-figure 3b; the first segment is short and 
rather broader than the second; the coalesced third to sixth segments form 
a distally narrowed piece and the terminal segment is triangular. The 
anterior male pleopod is flattened distally and its apex bends backwards. 

The chelipeds are not robust and are shorter than the walking legs; the 
propodus is rather heavy and the immovable finger is downturned proxi- 
mally on the prehensile margin. The ambulatory legs are rather stout; 
the first pair is a little shorter than the second and third pairs, and the 
last pair is the smallest. Each dactylus is flat and inwardly curved, the 
inner border is entire. 

Habitat: This crab is a commensal with a holothurian, Protankyra 
hidentata (Woodward and Barrett), which is very common on the mud 
flat of Shimabara, Nagasaki Prefecture. 

Measurements: Length of carapace 2.5 mm, width of same 3.5 mm. 



Family Majidae Alcock 
Genus Epialtus H. Milne-Edwards 

No species of the genus Epialtus has as yet been recorded from Japan 
and the adjacent waters. The majority of species are restricted to the 
east and west coasts of North, Middle and South America. The new 
Japanese species described below is the first recorded occurrence of this 
genus in the Eastern Indo-Pacific. 



252 Proceedings of the Biological Society of Washington 




Text-fig, 4, a. Epialtus orientalis new species. Dorsal view of male 
holotype. b. Sphenocarcinus bidens new species. Carapace in dorsal 
view. c. Chlorinoides tosaensis new species. Profile view of carapace of 
male holotype. (a X 3; b X 2.7; c X 1.8) 



Epialtus orientalis new species 
Text-fig. 4 a 

One $, holotype (SI 125883). Kii Shirahama. Sent by Dr. H. 
Uchinomi of the Seto Marine Biological Station. 

Diagnosis: This new species is closely related to Epialtus sulcirostris 
Stimpson from the Gulf of California, Mexico. The carapace is elongate 
pentagonal in outline. The rostrum is triangular with its apex pro- 
longed into a process directed obliquely downwards and its lateral 
borders set with hooked hairs. The preorbital tooth is acuminate, while 
the external orbital tooth is rudimentary, and the supra-orbital margin en- 
tire. 

The anterior half of the anterolateral border is anteriorly convergent 
and unarmed, while the posterior half bears three teeth, of which the 



New Japanese crabs 253 

middle one is the smallest and set a little closer to the third. The poster- 
olateral borders are slightly convex; the posterior border is very narrow 
with an obscure median tubercle. The dorsal surface of the carapace is 
almost flat, but the gastric and cardiac regions are somewhat convex. 

The chelipeds are a little stouter and longer than the ambulatory legs. 
The merus and propodus are subequal in length and thickness, while the 
carpus is short and its outer edge slightly cristate; the fingers are short 
and unarmed. The ambulatory legs gradually decrease in length from 
first to last. The anterior border of the merus has two lobules, that of the 
carpus only a large one, and that of the propodus two smaller ones. The 
dactylus of all pairs is hooked at the tip. 

The abdomen of the male holotype is composed of six segments, there- 
fore this species is referable to Epialtus, not to Epialtoides Garth 1968. 

Measurements: Length of carapace 9.1 mm, width of same 7.2 mm. 

Genus Sphenocarcinus A. Milne-Edwards 

Sphenocarcinus bidens new species 

Text-fig. 4 b 

Carapace only, sex unknown, holotype (SI 125884). Off Kumano- 
nada, Mie Prefecture; depth 150-180 m. Sent by H. Kato of the 
Aichi-ken Fisheries Experimental Station. 

Diagnosis: This new species is closely related to S. carbunculus Rath- 
bun from the Hawaiian Islands in the arrangement of the protuberances 
of the carapace, and also to S. stimpsoni (Miers) from Japan to Colombo, 
in the form of the process at the lateral angle of the carapace. 

The carapace is in the form of a rounded triangle, the rostral horns are 
divergent in the distal third, directed obliquely downwards. Their outer 
margins are subparallel. 

On the dorsal surface of the carapace, there are 13 raised protuberances 
in all, of which three are located in the median line, one each on the gas- 
tric, cardiac and intestinal regions — the gastric one is the largest and is 
longitudinally oval, the cardiac one is a rounded triangle with its base 
directed forward and the intestinal one is transversely narrow with its 
middle portion constricted. The rest of the protuberances are symmetri- 
cally disposed — one on the supra-orbital margin ( the smallest of all ) ; one 
on the hepatic margin is narrow and invaginated in the middle; three on 
the branchial region, of which two are large and located on the dorsal sur- 
face, the other one at the lateral angle is developed into a large obtuse 
process, the tip of which projects obliquely forward. Besides these pro- 
tuberances, there are a few tiny tubercles — one in front and one or two on 
the lateral sides of the gastric protuberance, and also one on the antero- 
lateral border between the hepatic and branchial protuberances. There are 
no tubercles or protuberances on the ventral side of the carapace but a 
few small granules on the ridge between the subhepatic and pterygostom- 
ian regions. 



254 Proceedings of the Biological Society of Washington 




Text-fig. 5. Anterior male pleopod of: a. Chlorinoides tosaensis new 
species, b, b'. Maja nagashimaensis new species. ( a X 20; b X 8; b' X 
50) 



Although the abdomen and thoracic legs are missing, the species may 
be safely designated as new to science. 

Measurements: Length of carapace 13.2 mm, width of same 9.0 mm, 
length of rostral horn 4.0 mm. 

Genus Chlorinoides Haswell 

Chlorinoides tosaensis new species 

PI. I, Fig. 3; Text-figs. 4 c; 5 a 

Two ^ ^ , 1 2 . One male ( SI 125882 ) is designated as the holo- 
type. Off Mimase, Tosa Bay. Coll. by K. Sakai. 

Diagnosis: This new species is closely related to Ch. acanthonotus 
Adams and White from Borneo, but the branchial regions are apparently 
more swollen and the rostral horns much longer, their total length being 
more than half that of the carapace. 



New Japanese crabs 255 

As in Ch. acanthonotus, the supra-orbital spine is extremely long and 
curved posteriorly in the distal half, but unlike that species, it is simple 
and not bifurcate in the distal half. A long and simple spine in the mid- 
dle portion of the intestinal region projects obliquely backward; in Ch. 
acanthonotus, however, it is divided into two proximally, one projecting 
upwards and the other horizontally backwards. The gastric region is 
armed with two long spines, one behind the other in the median line; the 
cardiac region also has two, side by side, set near to each other and di- 
verging somewhat. One hepatic spine of medium size, projects obliquely 
backward and finally two very long branchial ones near the antero-exter- 
nal corner, project obliquely forward in one case and obliquely backward 
in the other. 

The merus of the ambulatory legs is armed with two long spines at 
their distal extremity. In other respects the new species agrees well with 
C. acanthonotus. 

Measurements: Length of carapace of male holotype, 21 mm, width of 
same 17 mm, length of rostral horn 17 mm. 

Genus Leptomithrax Miers 

In the fauna of Japan, the genus Leptomithrax has hitherto been rep- 
resented by only two early recorded species — L. edwardsi de Haan and L. 
hifidus Ortmann. Each of these have the chelipeds rather robust and the 
carpus very short, unlike the typical species of New Zealand and Austra- 
lia, in which the chelipeds are slender and the carpus is relatively long and 
cylindrical. 

The new species described below has the chelipeds very long and 
slender, and the carpus is apparently longer as seen in the species of the 
southern hemisphere. 

Leptomithrax kiiensis new species 
PI. I, fig. 4 

One $ , holotype ( SI 125885 ) . Kii Minabe, from a lobster net in 
shallow water. Coll. by the late M. Ozaki. 

Diagnosis: The carapace is elongate pyriform and its outline, excluding 
the orbital and frontal portions, is narrower than in any of the Australian 
species. The dorsal surface is covered with a soft tomentum and numer- 
ous spinules. The spinules vary in size; the two gastric spines are on the 
median line, while the two cardiac ones are side by side; the two or three 
on the metabranchial region are most prominent. 

The rostral horns are slender and long, being more than half the length 
of the carapace proper. The supra-orbital spine is small and the inter- 
calated one moderate in size, but the external orbital one is very prominent 
and acuminate at the tip. On the anterolateral borders, there are four 
marginal spines — one hepatic and the others branchial. 

The chelipeds are very slender and more than twice as long as the 
carapace proper. The merus and carpus are cylindrical and beset with 



256 Proceedings of the Biological Society of Washington 

spinules, the latter about three fourths the length of the former. The prop- 
odus is markedly longer than the merus, smooth and glabrous. On the 
left cheliped, the propodus has a nodule on the upper and lower borders; 
on the right propodus, only the upper border has a nodule. Whether these 
nodules are due to an abnormality or are specific is not certain at present. 
The lower proximal border of the immovable finger has a large nodule near 
the proximal portion. The ambulatory legs are thickly covered with 
hooked hairs of almost equal length. 

Measurements: Length of carapace 55 mm, that of rostral spine 24 mm, 
width of carapace 44.5 mm, length of cheliped 125 mm. 

Genus Mafa Lamarck 
Maja nagashimaensis new species 

PL I, fig. 2; text-fig. 5b, b' 

Three $ $, 2 9 2. One male (SI 125886) is designated as the 
holotype: Kii Nagashima, Mie Prefecture. Coll. by Y. Yama- 
moto of the Nagashima High School. 

Diagnosis: In the general outline of the carapace, this new species is 
allied to M. japonica Rathbun and M. gibba Alcock, but in the number 
and arrangement of the spines on and around the carapace, it is related to 
M. miersi Walker. 

The carapace is markedly constricted behind the hepatic regions, and 
the branchial regions are well-inflated and round in outline. The dorsal 
surface is thickly covered with granules of various sizes. In the median 
line, there are two low spines, one on the gastric region and the other 
on the cardiac. The rostral spines are rather slender, short, well divergent 
apically and horizontal, not upturned as in M. miersi Walker or M. spini- 
gera de Haan. The extra-orbital spines are very strong and their anterior 
edge is excavated proximally. The hepatic spine is basally confluent with 
the external orbital one and its tip directed obliquely backward. The four 
marginal spines of the branchial regions are very strong and subequal, the 
final one is located on the dorsal surface, as in other species of this genus. 
The two usual spinules of the posterior border are almost rudimentary. 
The palp of the antenna arises from within the orbit. 

The chelipeds are robust, the merus short and distally thickened, colored 
with two or three irregular bands of deep purple. The carpus is robust and 
bears two longitudinal ridges, one median and the other on the outer 
margin; the propodus is strikingly swollen and smooth; the fingers 
are unarmed. The ambulatory legs are only sparsely hairy. The merus, 
carpus and propodus are also banded with deep purple. 

The anterior pleopod of the male is armed with a curved booklet near 
the apex as shown in text-figs. 5 b, b'. 

Measurements: Length of carapace proper 33.5 mm, width of same 
28.3 mm, length of rostral spine 8 mm. 

Relationship: Most of the species of Maja have the chelipeds very 
slender in both sexes, the carpus being relatively long and cylindrical; in 



New Japanese crabs 



257 




Text-fig. 6. Parthenope (Pseudolambrus) ozakii new species. Dorsal 
view of male holotype. ( X 3.4 ) 



the new species, however, the carpus is short and robust and the prop- 
odus also inflated. As regards the chelipeds, the new species thus ap- 
proaches some species of Leptomithrax, such as L. edwardsi de Haan and 
L. bifidus Ortmann, while in the form of the antenna it belongs to Maja, 
the antennal flagellum arising from within the orbit. In Leptomithrax, the 
flagellum is excluded from the orbit. 



Family Parthenopidae Alock 

Genus Parthenope Weber 

Subgenus Pseudolambrus Paulson 

Parthenope (Pseudolambrus) ozakii new species 

Text-fig. 6 

One ^, holotype (SI 125887). Kii Minabe, Wakayama Prefecture, 
shallow water. Coll. by the late M. Ozaki. 

Diagnosis: A rather small species; its nearest relative is probably P. 
( Ps. ) bicornis ( Flipse, 1936 ) from the Sea of Java, from which it may be 
distinguished by the peculiar lobe on the outer border of the propodus of 
the cheliped and by the different arrangement of the teeth and processes 
of the carapace. 

The carapace is in the form of a rounded triangle, the dorsal surface 
being smooth. The gastric and cardiac regions are markedly convex and 
conical, each ending in a long process, of which the gastric is thick and 



258 Proceedings of the Biological Society of Washington 

very long, projecting obliquely backwards, while the cardiac is a little 
smaller and projecting upwards. 

The front is rounded triangular in outline, the margin entire. On the 
anterolateral borders, the hepatic portion is entire and rounded, com- 
pletely merged with the external orbital angle; the branchial portion is set 
with 6-7 teeth, which are triangular and confluent with each other at the 
base. The carapace is very slightly constricted behind the hepatic margin. 
There is an oblique and sparingly granulated ridge on each branchial 
region running from the last anterolateral tooth toward the middle portion 
of the carapace. The posterolateral borders are almost straight and marked 
with a few tubercles. 

The chelipeds are strikingly asymmetrical. The left is very small, each 
segment is slender and the fingers cross each other strikingly in the distal 
half. The right cheliped is robust, the merus very broad and its anterior 
and posterior borders cut into five or six cristate teeth, the carpus is thin 
and small, while the propodus is again robust, its outer border bears a very 
broad and round lobe at the proximal half, its interior border is finely 
serrated. The immovable finger has three obtuse teeth near the middle 
of the prehensile edge. 

The ambulatory legs are very thin and the anterior and posterior borders 
of each segment are marked with tiny denticles. 

Measurements: Length of carapace 12.1 mm, width of same 12.5 mm. 

Family Cancridae Alcock 

Genus Cancer Linne 

Cancer nadaensis new species 

Text-fig. 7 

One $, holotype (SI 125871). Shimogusui, Kii Province, muddy 
bottom, depth 30 m. Coll. by H. Hayashi of the Odawara Car- 
cinological Museum. 

Diagnosis: The new species is distinguished from any other of the 
Japanese species of this genus by its well-cut anterolateral teeth, and also 
by the sharply and uniformly spinulated carpus and propodus of the 
chelipeds. The carapace is about 1.5 times as broad as long and its dorsal 
surface is slightly areolated, with a few obscure scattered tubercles near 
the anterior and lateral surfaces. The front is cut into three teeth, the 
middle one of which is thin and on a lower level. The preorbital tooth is 
prominent and sharp. 

The anterolateral borders are well arched and divided into nine teeth. 
All of the teeth are well-cut and separated from each other, their tips being 
acuminate and the last one largest of all and projecting sideways. On the 
posterolateral border, there is a small tooth just behind the last anter- 
olateral tooth, followed by a series of tiny teeth. 

The chelipeds are symmetrical. The merus is short and scarcely visible 
from above; the carpus and propodus are rather thickly and uniformly 



New Japanese crabs 



259 




Text-fig. 7. Cancer nadaensis new species. Dorsal view of male holo- 
type. (Xl) 

spinulated on the upper and outer surfaces, the spinules variable in size 
and irregularly arranged. 

The anterior three pairs of ambulatory legs are subequal in size, the last 
pair being a little smaller. The anterior margins of the merus, carpus and 
propodus are obscurely spinulated. 

Measurements: Length of carapace 32 mm, width of same 48 mm. 

Family Portunidae Dana 
Genus Benthochascon Alcock 

In the Indo-Pacific, the genus Benthochascon has been represented by 
a single species, B. hemmingi Alcock, which was originally recorded from 
the Andaman Sea. The second locality of this species is the Nicobars and 
the third Tosa Bay, Japan. A new species of this genus has recently been 
obtained at Izu Niijima, and is described below. 



Benthochascon elongatum new species 
Text-fig. 8 a, b 

One $, holotype (SI 125869), dried, ambulatory legs missing. 
Izu-Niijima, one of the seven islands of Izu. Sent by T. Oda- 
wara of the Odawara Carcinological Museum, Tokyo. 

One $ (photo), same locality. Sent by K. Suzuki of the Kanazawa 
Aquarium, Ishikawa Prefecture. 

Diagnosis: The carapace is oblong, almost 1.2 times as long as broad. 
The dorsal surface is smooth and glabrous. The front is composed of three 



260 Proceedings of the Biological Society of Washington 




Text-fig. 8. Benthochascon elongatum new species, a. Carapace of 
male holotype. b. Right cheliped of same, (a, b X 6) 



obtuse teeth, of which the median one is smaller than the lateral ones. 
Neither preorbital tooth or upper orbital fissures are present. The anter- 
olateral borders are cut into four teeth, the first or external orbital tooth is 
very long, the second, third and fourth are smaller and subequal in size. 
The posterolateral borders are much longer than the anterolateral ones and 
slightly concave. 

The basal joint of the antenna is short but longer than broad and freely 
movable; the flagellum is moderately long, situated in a narrow orbital 
hiatus. 

The chelipeds are rather robust; the merus short and prismatic, the 
carpus subquadrate in outline, armed with a tooth at the inner angle. The 
propodus is high, the immovable finger very short and proximally very 
broad, the outline being triangular. The movable finger is long and 
slender, strongly curved inward in the distal half. 

Measurements: Holotype, length of carapace 6.8 mm, width of same 
5.5 mm. 

Genus Thalamita Latreille 

Thalamita yoronensis new species 

Text-fig. 9 a, b 

One $, holotype (SI 125870), 1 2, allotype. Yoron Island, 

Amami Group. Coll. by K. Sakai. 
One ^,12, same locality. Coll. by Hirosuke Yamada. 

Diagnosis: Related to Thalamita picta Stimpson from. Japan to tropical 
Indo-Pacific and Th. macropus Montgomery from Abrolhos Islands, but 
the new species may be distinguished from them by the different features 
of the frontal, preorbital and anterolateral teeth. 

The carapace is little more than two-thirds as long as broad. The dorsal 
surface is covered with thin hairs and mottled with eight brown color pat- 



New' Japanese crabs 



261 




Text-fig. 9. Thalamita yoronensis new species, 
holotype. b. Abdomen of same. ( a X 3.5; b X 7 ) 



a. Carapace of male 



terns in the fresh specimen. There are two pairs of transverse ridges on the 
gastric region, one each on the protogastric and mesogastric. Behind the 
usual transverse ridge connecting the last anterolateral tooth on either side, 
is a short obscure ridge which traverses the cardiac region. 

There are 6 frontal teeth, the median 2 are close together, rounded at 
the tip and well-produced forwards; the submedian ones are well-sep- 
arated and short but also rounded at the tip; the lateral ones are again 
projected forward and their tips rounded. The pre-orbital edge is armed 
with a sharp tooth, which projects forward, not sideways as in other con- 
geners; below this tooth an acuminate tooth at the inner angle of the 
inferior orbital margin is seen in dorsal view. 

The five anterolateral teeth are uniformly acuminate — the first and 
second projecting forward, the third and fourth obliquely forward, of 
which the fourth is not distinctively reduced in size; the fifth is prominent, 
projecting strongly sideways. The posterior border is very narrow. 

The chelipeds are slightly asymmetrical; the merus unarmed, the carpus 
with its inner angle armed with a long sharp tooth and its outer extrem- 
ity with three spinules; the propodus is slightly swollen in the middle and 
its upper border has five teeth, of which one is proximal, two are on the 
inner margin and the rest are on the outer margin. The ambulatory legs 
are banded with dark green color; the posterior border of the propodus of 
the last pair is spinulated. 

The penultimate segment of the male abdomen is subquadrate in out- 
line, the lateral borders being parallel and obtusely angular at the antero- 
lateral angle. 

Measurements: Length of carapace 12.7 mm, width of same 19.0 mm. 



262 Proceedings of the Biological Society of Washington 







^^^Oo 



Text-fig. 10. Neoliomera richtersi (de Man) from Hawaii. Frontal 
and lateral portions of carapace, (x 1.5) 



Family Xanthidae Alcock 
Genus Neoliomera Odhner 

Following is a list of species of Neoliomera recorded up to the present: 

Neoliomera pub escens (H. Milne-Edwards, 1834) 

N. insularis (White, 1848) 

N. variolosa ( A. Milne-Edwards, 1873 ) 

iV. sundaica (de Man, 1888) 

N. richtersi (de Man, 1889) 

N. themisto (de Man, 1889) 

N. praetexta (Rathbun, 1900) 

N. sabaea ( Nobili, 1906 ) 

N. intermedia Odhner, 1925 

JV. nohilii Odhner, 1925 

iV. sakagutii Sakai, 1939 = Atergatis granulatus de Man, 1889 

( Cf . Forest & Guinot, 1961 ) 
N. striata Buitendijk, 1941 
N. ovata Tweedie, 1950 = Atergatis sp. (Cf. Forest & Guinot, 

1961) 
N. demani Forest & Guinot, 1961 
N. immigrans Edmondson, 1962 z= Atergatopsis sp. 

Forest & Guinot ( 1961 ) suggested that N. intermedia recorded by me 
from Japan ( 1939 ) was a synonym of N. pubescens, but I have now veri- 
fied the Japanese species based upon the form of the first pleopod of 
male, which is quite different from that of N. pubescens as shown in text- 
figure 12 d. 

According to the same authors, N. ovata Tweedie is not Neoliomera but 
belongs to Atergatis. In my opinion, N. immigrans Edmondson is also not 
Neoliomera but may be a species of Atergatopsis. 

The present opportunity is taken to add two new species to this genus, 
which are described below. 



New Japanese crabs 



263 




Text-fig. 11. a. Neoliomera acutidens new species. Dorsal view of 
male holotype. b. Neoliomera richteroides new species. Dorsal view of 
male holotype. (a X 1.2; b X 2.1) 



Neoliomera acutidens new species 
Text-figs. 11 a; 12 c. 

Two $ $ , one of which is designated as the holotype ( SI 125874 ) . 
Shimogusui, Kii Province, trawled from 30-50 m. Sent by H. 
Hayashi of the Odawara Carcinological Museum. 

Diagnosis: The nearest relatives of the new species are N. pubescens 
and N. intermedia, but the following particulars may be enumerated as 
the differences between these species. 



264 Proceedings of the Biological Society of Washington 

The carapace is transversely ovoid, the dorsal surface being vaulted on 
the anterior one-third, but almost flat on the posterior two-thirds. The 
anterior and lateral surfaces are covered with depressed granules, of 
which those near the anterolateral teeth are larger. The meso- and meta- 
gastric regions, as well as the cardiac, intestinal and inner sub-branchial 
surfaces are flat and almost smooth. The grooves which define the regions 
are narrow but deep. In N. pubescens, the entire dorsal surface is uni- 
formly covered with granules. 

The frontal margin is divided into two lobes by a median fissure, each 
lobe being almost transversely straight. The antero-lateral borders are cut 
into four teeth, of which the first slants outward and is completely con- 
fluent with the outer orbital angle; the second is a little shorter and low, 
while the third and fourth are triangular in shape, each having a pointed 
apex. In N. pubescens, as well as in N. intermedia, these four teeth are 
rounded and not at all dentate. 

The chelipeds of the male are subequal, the carpus is marked with a T- 
shaped groove in the anterior portion of the upper surface, and its inner 
angle is broadly produced, divided into obtuse teeth at the end. The prop- 
odus is laterally depressed. Its upper outer surface is longitudinally 
grooved and the lower surface is almost smooth. The black colour of the 
immovable finger extends onto the outer and inner surfaces of the prop- 
odus, but does not expand so as to encircle the whole segment. 

All pairs of ambulatory legs are very slender compared with those of 
the species referred to above; the anterior margin of each segment is 
sparingly granulated, but the surface of each segment is almost smooth. 

The anterior pleopod of the male is curved inward at the tip and 
marked with a small obtuse process, as shown in text-figure 12 c. 

Measurements: Length of carapace 21 mm, width of same 37.5 mm. 
Neoliomera richteroides new species 
Text-figs. 11 b; 12 b. 
Neoliomera richtersi Sakai 1967 (nee N. richtersi de Man), Re- 
searches on Crustacea, Tokyo, no. 3, p. 81, Frontispiece fig. 1 
( colored ) . 
One $ , holotype ( SI 125875 ) . Yoron Island, Amami Group, coral 
reef. Coll. by K. Sakai. 

Diagnosis: One male specimen obtained from the Yoron Island, which 
was formerly reported by the author as N. richtersi de Man (loc. cit. ) 
seems to be a distinct new species. The author examined the typical N. 
richtersi from Bikini Island by courtesy of Dr. J. S. Garth at the Allan 
Hancock Foundation, Los Angeles, and several male and female specimens 
from the Hawaiian Islands identified by Edmondson, by courtesy of Dr. 
D. M. Devaney at the Bemice P. Bishop Museum, Honolulu. 

The carapace is appreciably broader than long and the dorsal surface 
is thickly studded with granules of various sizes. The anterolateral borders 
are well arched and distinctly cristate, cut into four teeth, of which the 
first and second are long and lobular, obscurely divided by an indistinct 



New Japanese crabs 



265 




Text-fig. 12. Anterior pleopod of male of: a. Planopilumnus minahen- 
sis new species, b. Neolwmera richteroides new species, c. Neoliomera 
acutidens new species, d. Neoliomera intermedia Odhner from Kii Penin- 
sula, (a X 41; b X 72; c X 62; d X 62) 



incision, while the third and fourth are dentiform, the former as long as 
the third and the latter very short. In typical N. richtersi, the anterolateral 
borders are not arcuate but slender, the first and second teeth entirely 
united and the third and fourth are short. The margins of these teeth are 
studded with sharp granules and the crest is very narrow. 

The frontal and upper orbital margins are also distinctly cristate, while 
in N. richtersi, they are not cristate at all. 

The chelipeds resemble those of N. richtersi as does the black color of 
the immovable finger which extends onto the surface of the propodus, 
enclosing the distal portion of that segment. 



266 Proceedings of the Biological Society of Washington 




Text-fig. 13, Planopilumnus minabensis new species. Dorsal view of 
male holotype. (X 2) 

The coloration of the fresh specimen is yellow as in N. nchtersi, having 
a number of whitish spots of various sizes on the carapace. 

The anterior male pleopod also resembles that of N. richtersi, having 
eight long and two or three shorter hairs at the distal portion. 

Measurements: Length of carapace 12.5 nmi, width of same 24.0 mm. 



Genus Planopilumrms Balss 

Planopilumnus minabensis new species 

Text-figs. 12a; 13 

One $,2 $ $, one male, holotype (SI 125876). Kii Minabe, 
Wakayama Prefecture, from the refuse of lobster net. Coll. by 
the author himself. 

One $, I 9- Same locality. Coll. by the late M. Ozaki. 

One $, Shimogusui, Kii. Coll. by H. Hayashi of the Odawara 
Carcinological Museum, Tokyo. 

Diagnosis: The entire body is thickly covered by hairs, which, on the 
dorsal surface of the carapace and chelipeds have a vermiform appear- 
ance, and on the ambulatory legs, are very long and fringe both borders 
of each segment. On denudation, the dorsal stuface of the carapace is 
well-areolated by deep, smooth grooves. Each areola has a shallow de- 
pressed surface, which is surrounded by an obsciu-e ridge marked with 
tiny granules. The front protrudes well forward, being divided into two 
truncate lobes, the free edges of which are granulated. The upper orbital 
margin is thick and has two distinct fissures. Behind the frontal lobes 



New Japanese crabs 267 

there are two pairs of granulated ridges, the anterior one being transverse 
and the posterior one obHque. 

The anterolateral border is armed with four teeth, all acuminate and spar- 
ingly granulated; the last one is a little smaller than the rest. The postero- 
lateral borders are sub-equal to the anterolaterals, being straight and 
moderately convergent. The posterior border is slightly convex. 

The chelipeds are robust and symmetrical; the merus is short but dis- 
tally thickened and armed with a subterminal tooth on the upper ridge. 
The carpus is large and swollen, with a shallow dorsal and an oblique 
subdistal groove. The propodus is rather short but high with a dorsal 
groove near the superior edge, the outer surface has four longitudinal 
rows of granules, but the inferior surface is uniformly granulated. Both 
fingers are lightly pigmented, the pigment does not extend onto the sur- 
face of the propodus. The ambulatory legs are robust, thickly fringed 
with hairs. On denudation, the carpus has a deep groove on the upper 
surface and the propodus has a shallow groove on each upper and lower 
surface. 

The abdomen is seven-segmented, the first and third terga are broad- 
est and the terminal tergum is rounded triangular. The anterior pleopod 
of the male is curved in the distal portion, as figured in text-figure 12 a. 

The new species is obviously related to P. vermiculatus ( A. Milne-Ed- 
wards) from New Caledonia and also to P. labyrinthicus (Miers) from 
north and north-east Australia. However, the vermiform sculptures of the 
carapace of the new species is quite different and the hair on the carpus 
and propodus also assumes a vermiform appearance, while in the above 
species, these segments are simply haired. 

The holotype of the new species was compared with that of P. vermi- 
culatus by courtesy of Forest and Guinot at Museum National d' Histoire 
Naturelle, Paris. 

Measurements: Holotype, length of carapace 14.8 mm, width of same 
20.5 mm. 

Genus Heteropilumnus de Man 

Heteropilumnus mikawaensis new species 

Text-fig. 14 a-c 

One $, holotype (SI 125873). Off Mikawa, Aichi Prefecture, 
depth 100-150 m. Coll. by H. Hayashi of the Odawara Car- 
cinological Museum, Tokyo. 

Diagnosis: The entire animal is sparingly covered with long hairs; the 
carapace is laterally subquadrate, its dorsal surface being vaulted on the 
anterior surface and almost flat from side to side in the middle and poste- 
rior surfaces. The regions are only obscurely defined. The front consists 
of two lobes, fringed with long hairs. 

The anterolateral borders are cut into four teeth, of which the first is 
entirely confluent with the external orbital angle and its free edge is 



268 Proceedings of the Biological Society of Washington 




.^/N-0/O/C\;jV,,s 






y 



^v-' 



Text-fig. 14. Heteropilumnus mikawaensis new species, a. Carapace of 
male holotype viewed from dorsal side. b. Anterior male pleopod. c. 
Chelipeds viewed from outer side, (a X 3.2; b X 70; c X 2.7) 



obliquely cut, the second and third are obtuse and well-separated, the 
fourth is very small and acuminate. These four teeth are granulated when 
viewed under the microscope. 

The left cheliped is small and the carpus, propodus and the proximal 
portion of the dactylus are studded with tubercles; the left cheliped is 
much larger than the right, the propodus being very heavy and the outer 
proximal surface with a few tiny scattered tubercles; otherwise the entire 
surface is smooth. 

The ambulatory legs are very slender, each segment being flattened. 
Both anterior and posterior borders are thickly fringed with hair. 

The male abdomen is seven-segmented, the first and third segments 
very broad as usual, and the fourth to seventh very narrow. The first 
male pleopod is looped near the apex and its inner border is beset with 
several setae, one of which is strikingly thick and long as shown in text- 
figure 14 b. 

Measurements: Length of carapace 9.8 mm, width of same 13.6 mm. 

Relationship: This new species appears to be related to H. trichophorus 
(de Man, 1895) from Malakka, but the entire body is not tomentose and 
the carapace is fairly well areolated; the anterolateral teeth are, unlike that 
species, covered with granules. The ambulatory legs are much more 
slender. 



New Japanese crabs 



269 




f 

Text-fig. 15. Six species of Carcinoplax from Philippines: a. C. bi- 
spinosa Rathbun. Type male. b. C. angusta Rathbun. Type female, c. C. 
specularis Rathbun. Type male. d. C. verdensis Rathbun. Type female, 
e. C. confragosa Rathbun. Type female, f. C. spinosissima Rathbun. Type 
male, (a X 1.3; b X -8; c X .9; d X 1.5; e X .5; f X .7) 



Family Goneplacidae Dana 
Genus Carcinoplax H. Milne-Edwards 

At present, the Indo-Pacific genus Carcinoplax contains about 16 spe- 
cies including one subspecies. Some of these species are insufficiently rec- 
ognized and have not yet been illustrated. In my opinion, these species 
are divided into three groups on the basis of the form of the orbital and 
anterolateral teeth. 

In the first or typical group, which has the carapace rather vaulted 
fore and aft, the anterolateral borders are armed with three teeth, the first 
of which arises at the external orbital angle, and the second and third 
are rather dentiform, having the tips sharply pointed. In some species, 
however, these teeth are obliterated in the fully grown stage. 

The following species belong to this group : 

Carcinoplax longimanus ( de Haan, 1835 ) 

C. vestitus (de Haan, 1835) 

C. eburnea Stimpson, 1858 

C. longimanus indicus Doflein, 1904 

C. purpurea Rathbun, 1914 

C. bispinosa Rathbun, 1914 

C. angusta Rathbun, 1914 

C. meridionaris Rathbun, 1923 

C. victoriensis Rathbun, 1923 

C. surugensis Rathbun, 1932 

C. inaequalis (Yokoya, 1933) 



270 Proceedings of the Biological Society of Washington 




Text-fig. 16. a. Carcinoplax tomentosa new species. Dorsal view of 
male holotype, (x 1.4) b. Psopheticoides sanguineus new genus, new 
species. Dorsal view of male holotype. (x 1.2) 



In the second group, which has the carapace rather depressed, the 
anterolateral borders are armed with three dentiform teeth, the first of 
which is obtuse or transverse in form, and is clearly separated from the 
external orbital margin by a connecting ridge. 

The following species belong to this group : 

Carcinoplax longipes (Wood-Mason, 1891) 
C. verdensis Rathbun, 1914 
C. specularis Rathbun, 1914 
C. tomentosa new species 

In the third group, which has the carapace vaulted fore and aft, and 
the entire body tomentose and granulated, the anterolateral borders are 
armed with four spinulated teeth, the first of which is the external orbital 
one. This group seems to be aberrant and should possibly be separated 
from the typical Carcinoplax. 



Netv Japanese crabs 271 



The following species belong to this group : 

Carcinoplax confragosa Rathbun, 1914 
C. spinosissima Rathbun, 1914 



By permission of the Smithsonian Institution, the types of six Philippine 
species of Carcinoplax described by Rathbun are illustrated for the first 
time in text-fig. 15. 

Carcinoplax tomentosa new species 
Text-figs. 16a; 17c; 18a. 

One ^, holotype (SI 125872). Off Mimase, Tosa Bay, depth 150- 

200 m. Coll. by K. Sakai. 
Three 2 $ . Same locality, depth 100 m. Same collector. 
One $ (photo). Osaka Bay. Coll. by H. Kohno of the Osaka 

Municipal Museum. 

Diagnosis: A medium-sized crab. The entire animal is covered with 
thin, soft hairs. The carapace is sensibly broader than long, the sur- 
face depressed and the regions scarcely delimited. The front protrudes 
forward beyond the level of the preorbital angles and its free edge is 
truncate, the median fissure being obscure. The upper orbital margin 
is transverse in its outer half and directed rather obliquely forward. 

The anterolateral border is cut into three teeth; the first tooth is broadly 
truncate and independent, being separated from the outer edge of the 
orbital margin by a connecting ridge; the second and third teeth are 
pointed with their tips directed forwards. The posterolateral margins are 
a little longer than the anterolateral, and are slightly convergent back- 
wards. 

Chelipeds are heavy and almost symmetrical; the merus is rather thin 
and prismatic, with a tiny process near the distal end of the upper inner 
ridge; the carpus is rhomboidal in outline with its inner angle armed with 
a strong tooth, and its outer border armed with a tiny spine; the propodus 
is inflated on the outer surface and its upper inner border is obscurely 
cristate; the fingers are nearly as long as the propodus, their distal half 
being pigmented and their prehensile edges sharply denticulated. 

The anterior three pairs of ambulatory legs are subequal in length and 
their dactyli are slender, slightly depressed and lanceolated. The last 
pair of legs are distinctly shorter and the propodus and dactylus are 
broader than in the preceding pairs. The male abdomen is triangular in 
outline and the terminal segment is longer than any of the other segments. 

Measurements: Length of carapace 24.7 mm, width of same 31.5 mm. 

Relationship: In the form of the front and anterolateral teeth, the new 
species appears to be closely related to C. specularis Rathbun, but the en- 
tire body is tomentose and the anterior pleopod of the male is spatulated 
and obtusely projecting at the tip; in C. specularis, the distal portion of 
the same appendage is obtusely bilobate. 



272 Proceedings of the Biological Society of Washington 




Text-fig, 17. a, a'. Anterior pleopod of male of Psopheticoides sanguin- 
eus new genus, new species b. Posterior pleopod of same. c. Anterior 
pleopod of male of Carcinoplax tomentosa new species, (a X 13; a' X 
52; b X 13; c X 12) 



Genus Psopheticoides new genus 
Type-species: Psopheticoides sanguineus new species. 

This new genus is allied to Carcinoplax H. Milne-Edwards, Psopheticus 
Wood-Mason and Eucrate de Haan. 



New Japanese crabs 



273 




Text-fig. 18. a. Anterior part of carapace of Carcinoplax tomentosa 
new species, b. Same of Psopheticoides sanguineus new genus, new spe- 
cies, c. Male abdomen of Psopheticoides sanguineus new genus, new spe- 
cies, (a X 1.4; b X 2.1; c X 3.8) 



In its general features, the carapace of the new genus is related to 
Carcinoplax, but the dorsal surface is depressed and the anterolateral 
borders are more strongly arched and cut into three cristate, lobular teeth, 
the anterior two of which are sometimes confluent. The posterolateral 
borders are markedly convergent backwards and concave. 

The eyes and orbits are peculiar and resemble those of Psopheticus. The 
eye-stalks are short and thick and the cornea very large and inflated. The 
orbits are very large and deep to accommodate the large cornea; the upper 
orbital margins are strikingly sinuous. 

The abdomen and pleopods are related to those of Eucrate, the male 
abdomen being very narrow and elongate in outline, composed of seven 
distinct segments. The anterior male pleopod is very long and thin, and its 
apex armed with a hook; the posterior one is very short and its tip 
slanting. In Carcinoplax and Psopheticus, the male abdomen is broadly 
triangular in outline and the anterior male pleopod is rather flat and 



274 Proceedings of the Biological Society of Washington 

spatulate at the tip; the posterior pleopod is as long as or longer than the 
first and its tip bifurcated. 

The dactylus of the 4th ambulatory legs is flat and somewhat lanceo- 
late, suggesting relationship with the genera of Catoptrinae. 

Psopheticoides sanguineus new species 
Text-figs. 16b; I7a; 18b,c 

One ^,12, the male (SI 125888) is designated as the holotype. 
Mimase, Tosa Bay, depth 100-150 m. Coll. by K. Sakai. 

Diagnosis: This new species is peculiar in having a round, deep red 
spot, surrounded by a white rim, in the center of the carapace. The 
carapace is anteriorly broad and the posterolateral borders convergent 
backwards. The frontal margin is truncate, about one-fourth the greatest 
width of the carapace; there is no median notch. The anterior end of the 
orbital eave has a small tooth, which is separated from the edge of the 
front by an indistinct fissure. 

The anterolateral borders are markedly arched, somewhat cristate and 
cut into three lobes ( in one female, the left side has only two lobes, the 
first and second ones being fused together). 

The chelipeds are heavy and symmetrical; the merus is rather short and 
its upper border obscurely ridged, with a subterminal tooth. The carpus is 
thick, as long as the merus, and has a sharp spiniform process near the 
middle of the inner border. The propodus is also very thick, the fingers are 
not pigmented, the prehensile edges are armed uniformly with small sharp 
teeth. The anterior three pairs of ambulatory legs are subequal in length 
and thickness. The last pair is a little shorter and the carpus distally broad, 
the propodus moderately broad and flattened and the dactylus somewhat 
lanceolate in form. The propodus and dactylus are haired along both 
margins. 

The abdomen of the male is slender, the 4th to 7th terga being narrow, 
and the general outline resembles that of Eucrate. The anterior male 
pleopod is very slender and its distal portion armed with a hook at the 
tip, and four tiny spinules on the outer margin. The posterior pleopod is 
very short and its distal portion slanting. 

Measurements: Length of carapace of male holotype, 20.5 mm, width 
of same 26 mm. 

Relationship : In general features of the body, the new species seems to 
be related to Pilumnoplax cooki Rathbun 1906 from Hawaii, but the latter 
species has the eyestalks slender and the cornea not at all inflated, and 
the dactylus of the last pair of ambulatory legs slender and not lanceolated. 

The Indian species, Nectopanope rhodobathes Wood-Mason also seems 
to be allied to the new species, but the carapace is not broadened an- 
teriorly and the eyestalks are slender. Also, the cornea is not markedly in- 
flated. 



New Japanese crabs 275 



Family Pinnotheeidae Dana 
Genus Orthotheres new genus 
Type-species: Orthotheres turhoe new species. 

Among numerous species of the genus Pinnotheres, some are unusual 
in having the dactylus of the external maxilliped very small and inserted 
at the end of the propodus. This is unlike the usual features of the genus, 
in which the dactylus is stiliform and inserted on the inner side of the 
propodus. In such unusual species, the carapace of the female is ap- 
preciably broader than long and the dactylus of all pairs of ambulatory 
legs uniformly very short and sharply hooked. 

Ecologically, such species are found in the stomach of gastropods, but 
some are found in bivalve moUusks as in the common species of Pin- 
notheres. 

In the classification of the genera of the Pinnotheridae, the shape of 
the dactylus and its position relative to the propodus of the external max- 
illiped are regarded as important criteria. The author takes the present 
opportunity to establish a new genus to accommodate such unusual species, 
separating them from the genus Pinnotheres, and designating the new 
Japanese species, Orthotheres turhoe as the type-species. 

At least, the following American species may be safely included in the 
new genus: 

Pinnotheres serrei Rathbun Host: Stromhus. 

P. harhatus Rathbun Host: Turbo. 

P. stromhi Rathbun Host: Stromhus. 

Among the Indo-Pacific species, the following may be included in the 
new genus, although the carapace is rather narrow and the host is either 
a bivalve moUusk or unknown. 

Pinnotheres longipes Biirger Host: unknown. 

P. laevis Biirger Host: Coralliophaga. 

Orthotheres turboe new species 
PI. II, figs. 1, 2; text-fig. 19a. 

Three ^ ^, 3 $ $, one female (SI 125889) is designated as the 
holotype. Yoron Island, Amami Group. Goll. by K. Sakai and 
H. Yamada. 

Description of female: The entire animal is soft, as in the usual species 
of Pinnotheres. The coloration in the living stage is light lemonish white. 
The carapace is very broad and transversely elliptical and anteriorly con- 
vergent; the ratio between length and breadth is 1 : 1.5. The dorsal sur- 
face is evenly convex and smooth; the front is declivous, its margin being 
slightly bilobate. The anterolateral borders are markedly divergent back- 
wards; the posterior margin is very broad, a little convex in the middle. 

The dactylus of the external maxillipeds is very small and slightly curved 
inward near the tip, inserted at the end of the propodus. 

The chelipeds are stout and longer than any of the ambulatory legs; the 



276 Proceedings of the Biological Society of Washington 




Explanation of Plate II 

Fig. 1. Orthotheres turboe new genus and species. Female holotype, 
dorsal views. X 3.5. 

Fig. 2. Orthotheres turboe new genus and species. Male allotype, dorsal 
view. X 3. 



New Japanese crabs 277 

propodus is distally thickened and the dactylus armed with a tooth near 
the proximal end of the prehensile edge. The movable finger is unarmed. 

The anterior three pairs of ambulatory legs are subequal in length; the 
last pair is slightly more slender and shorter than the preceding ones. The 
dactylus of all pairs is uniformly short and hooked at the tip. 

Measurements: Length of carapace 8.5 mm, width of same 13 mm. 

Host: Turbo (Marmarostoma) argyrostomus (Linne); found in the 
stomach. 

Description of male: The male is always living with the female in the 
stomach of the same host. The entire body is fairly well calcified. The 
carapace is narrower than in the female, rounded quadrangular in out- 
line. The ratio between length and breadth is approximately 1:1. The 
front protrudes well-beyond the level of the eyes, its margin being obtuse 
and obscurely bilobate. 

The chelipeds are robust and the propodus distally thickened; the mov- 
able finger is strikingly curved inward at the tip and armed with a tooth 
near the proximal portion of the prehensile edge. The movable finger is 
unarmed. The features of the ambulatory legs are almost the same as in 
the female. In the male of Pinnotheres, the ambulatory legs are usually 
haired along the anterior and posterior borders of the carpus, propodus 
and dactylus, but in the present case, all such segments are naked. 

Measurements: Length of carapace 5.0 mm, width of same 5.1 mm. 

Host: Same as the female. 

Remarks: The same crab was formerly described by K. Nakasone from 
the Palao Island (1937, Nanyo Suisan Jyoho, no. 1, p. 32), but no name 
was given by the writer. The host was also Turbo ( Marmarostoma ) argy- 
rostomus (Linne). 

Genus Sakaina Serene 

The genus Sakaina was established by Serene on the basis of S. japo- 
nica, which is related to S. asiatica (Sakai). The author takes the present 
opportunity to add a new species to this genus, which was obtained on the 
coast of Manazuru, Sagami Bay. 

Sakaina incisa new species 
Text-figs. 19b; 20 
Two $ $, 2 2 2, one male (SI 125878) is designated as the 
holotype. Manazuru, Sagami Bay, rocky shore. Coll. by the 
author. 
Diagnosis: A tiny species like other congeners. In general view the 
new species is closely related to S. japonica Serene and also to S. asiatica 

<r 

Fig. 3. Macrophthalmus (Macrophthalmus) ceratophorus new species. 
Male holotype: a. General view of carapace and thoracic legs in dorsal 
view. X 1.4. b. Ventral view of carapace showing the abdomen. X 1.2. 
c. Chelipeds viewed from outside. X 2.8. d. Left half of carapace showing 
the eyestalk with its palp. X 2.8. 
X 1. 



278 Proceedings of the Biological Society of Washington 




Text-fig. 19. a. External maxilliped of Orthotheres turboe new genus, 
new species, b. Male abdomen of Sakaina incisa new species, (a X 70; 
b X 20) 



( Sakai ) . The carapace is transversely quadrilateral. The front protrudes 
slightly beyond the eyes and is medially notched. Just behind the frontal 
lobes the anterior surface of the carapace is defined by a rim of pu- 
bescence. 

The anterior half of the anterolateral border is evenly rounded, with a 
thick rim of marginal pubescence which, however, unlike that of S. asiatic 
or S. japonica, does not reach the external orbital portion. The dorsal sur- 
face is smooth and glabrous. 

The chelipeds are symmetrical, the merus small, and the carpus swol- 
len with its upper inner angle very slightly produced. The propodus is 
also swollen and its upper inner edge is slightly cristate; the fingers are 
unarmed. 



New Japanese crabs 279 




Text-fig. 20. Sakaina incisa new species. Dorsal view of male holo- 
type. (X 7) 

The anterior two pairs of ambulatory legs are subequal in length and 
thickness; the third pair is shorter and more slender, and the last pair is 
strikingly reduced. The abdomen is peculiar in that the terminal segment 
of the male is oblong and incised at the distal end. In S. japonica it is 
truncate and in S. asiatica it is rounded. 

Measurements: Length of carapace 2.5 mm, width of same 4 mm. 

Family Ocypodidae Ortmann 
Genus Macrophthalmus Latreille 

In 1967, Barnes revised the species of the genus Macrophthalmus and 
divided it into six subgenera — Macrophthalmus, Mareotis, Mopsocarcinus, 
Venitus, Hemiplax and Tasmanoplax; the number of species comprised in 
these subgenera are 41 in all. Of these species, four (Macrophthalmus 
transversus (Latreille, 1817); M. telescopicus (Owen, 1839); M. latipes 
Borradaile, 1903; M. milloti Crosnier, 1965) are peculiar in having the 
eye-stalks particularly long and reaching far beyond the external orbital 
angle. A new species from Japan, described below, belongs to this small 
group and is unusual in having a segmented palp on the distal end of the 
long eye-stalk. 

With regard to the subgeneric status, the new species agrees most 
closely with the chacteristics of Macrophthalmus in having the anterior 
boundary of the buccal frame produced into a process in the median por- 
tion; the teeth of the lateral margin of the carapace are basally narrowed 
and pointed; and, finally, the male abdomen relatively broad. In some 
respects, however, the new species represents the characteristics of the 
subgenus Mareotis, viz., the carapace being narrower, the ratio between 
length and width approximately 1 : 1.5, and the branchial region marked 
with two longitudinal rows of granules, and finally, the outer surface of 
the propodus of the chelipeds marked with a ridge of granules along the 
inferior border. 

A patch of soft hair found on the inner surface of the propodus of the 
chelipeds suggests the relation of this species to the subgenus Hem^iplax. 



280 Proceedings of the Biological Society of Washington 

Macrophthalmus ( Macrophthalmus ) ceratophorus new species 
PL II, figs. 3a-d 

One S, holotype (SI 125879). Shimogusui, Kii Province. Coll. 

by H. Hayashi of the Odawara Carcinological Museum, Tokyo. 
One $ (photo). Gokashyo Bay, Mie Prefecture. Col. by S. Tanaka 

of the Board of Education, Mie Prefecture. 

Diagnosis: The carapace is quadrilateral, the lateral borders slightly 
convergent backwards. The dorsal surface is rather depressed; the gastric 
region is well-defined by a distinct groove on either side and its surface 
is glabrous and punctate. The branchial regions are declivous on the outer 
half and granulated; there are two longitudinal rows of granules sub- 
parallel to the lateral border. The front is strongly constricted between 
the bases of the eye-stalks and its free edge is bilobed and deflexed. The 
external orbital tooth is not strong but sharp and is followed by two in- 
distinct teeth; otherwise the lateral border is unarmed. 

The anterior boundary of the buccal frame is clearly produced into an 
obtuse process, which is a principal criterion of the subgenus Macrophthal- 
mus as defined by Barnes. 

The eye-stalks are peculiar to this species, being much longer than the 
carapace and their tips reaching far beyond the external orbital angle. At 
the distal end, there is a thin palp composed of eight segments. 

The chelipeds are also peculiar to this species; the merus is slender and 
its anterior border finely granulated and sparingly setose; the carpus is 
small, its inner angle being armed with a spine and its outer border armed 
with several small obtuse spines; its inner surface is granulated and fur- 
nished with a patch of soft tomentum near the distal portion. The im- 
movable finger is unarmed and its prehensile edge almost entire, while the 
movable finger bears five long spines on the superior margin and a large 
tooth near the proximal end of its prehensile edge. 

The male abdomen is rather wide, the penultimate segment having an 
obtuse tubercle on each outer border; the terminal segment is rounded at 
its distal end. The anterior male pleopod has an apical process furnished 
with long hairs. 

Measurements: Length of carapace 19.5 mm, width of same 31 mm. 

Literature Cited 

Barnes, R. S. K. 1967. The Macrophthalminae of Australasia; with a re- 
view of the evolution and morphological diversity of the type 
genus Macrophthalmus. Trans. Zool. Soc. London, 31: 195- 
262. 

Forest, J., et D. Guinot. 1961. Crustaces decapodes brachyoures de 
Tahiti et de Tuamotu. Expedition frangaise sur les recifs coral- 
liens de Nouvelle-Caledonie, volume preliminaire : 80-86. 

Rathbun, M. J. 1914. A new genus and some new species of crabs of the 
family Goneplacidae. Proc. U.S. Nat. Mus., 48 (2067): 137- 
154. 



r 



Vol 



QH 

i 
B4X 



3 October 1969 



PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 

NEW CRAYFISHES FROM GEORGIA/^^ ^'^^ 
( DECAPODA ASTACIDAE ) f nfsT ^ 0^ a^^ 

' OCT 20 1969 

By Horton H. Hobbs, Jr. and Edward T. Hal^SiJr.^ /^^^^^j%,c - 

Smithsonian Institution and Georgia Water ^^*^- — "^ 
Quality Control Board 

Among the several undescribed crayfishes of the genus Cam- 
harus occurring in the Piedmont and submontane provinces of 
Georgia are two species, one which frequents riffles in the 
Chattahoochee Drainage System and the other which inhabits 
several clear, swiftly flowing tributaries to the Tennessee River 
in the northwestern corner of the State. 

One of them is here assigned to the typical subgenus which 
ranges from northern Georgia to New Brunswick and westward 
to Indiana, and the other to the subgenus Depressicambarus 
which is restricted to the southeastern portion of the United 
States, from North Carolina and Kentucky southward to the 
Florida panhandle and southern Mississippi. ( See Hobbs, "in 
press" for diagnosis of the subgenera. ) 

Cambarus (Cambarus) howardi, new species 

Diagnosis: Body pigmented, eyes well-developed. Rostrum with 
strongly convergent margins devoid of marginal spines or tubercles. Areola 
3.0 to 4.9 times longer than broad, comprising 34.3 to 39.3 percent of en- 
tire length of carapace, and bearing 3 to 6 punctations across narrowest 
part. Lateral surface of carapace with small cervical tubercles but lacking 
cervical spines. Suborbital angle obtuse. Postorbital ridges short with 
rounded or acute cephalic tubercles. Antennal scale approximately 2.3 
times longer than broad. Chela with single row of tubercles on mesial 
margin of palm; lateral margin of hand costate distally for at least half its 
length and both fingers with well-defined longitudinal ridge on upper sur- 
faces. First pleopod (Figs. 2, 4, 6) of first form male with corneous cen- 
tral projection recurved at approximately 110°, not markedly tapering dis- 
tally, with distinct subterminal notch, and not extending so far caudally as 
tip of mesial process; mesial process inflated, directed caudolaterally and 

21— Proc. Biol. Soc. Wash., Vol. 82, 1969 (281) 



282 Proceedings of the Biological Society of Washington 




Figs. 1-12. Cambarus howardi new species ( pubescence removed from 
all structures illustrated except for Fig. 2). 1, Lateral view of carapace 
of holotype. 2, Mesial view of first pleopod of holotype. 3, Mesial view 
of first pleopod of morphotype. 4, Caudal view of first pair of pleopods of 
holotype. 5, Lateral view of first pleopod of morphotype. 6, Lateral view 
of first pleopod of holotype. 7, Epistome of holotype. 8, Bases of third, 
fourth, and fifth pereiopods of holotype. 9, Antennal scale of holotype. 
10, Dorsal view of carapace of holotype. 11, Annulus ventralis and ad- 
jacent sternal elements of allotype. 12, Dorsal view of distal podomeres 
of cheliped of holotype 



New Georgia crayfish 283 

Table 1. Measurements (mm) of Cambarus howardi. 











Holotype 


AUotype 


Morphotype 


Carapace: 














Height 








10.1 


12.1 


8.3 


Width 








15.0 


17.0 


11.6 


Length 








27.0 


32.0 


22.6 


Areola: 














Width 








2.2 


2.6 


2.1 


Length 








9.9 


12.2 


8.0 


Rostrum: 














Width 








4.4 


5.2 


3.7 


Length 








6.3 


5.6 


4.6 


Chela: 














Length of 


inner 


margin 


of palm 


8.6 


8.7 


5.7 


Width of palm 






13.0 


12.9 


8.4 


Length of 


outer 


margin 


of hand 


25.4 


26.6 


16.5 


Length of dactyl 






15.3 


16.0 


10.5 



tip usually bi- or trispinose. Annulus ventralis (Fig. 11) concave with 
elevated lateral portions, with conspicuous tongue extending under dex- 
tral or sinistral wall. Color olive to light green with scarlet markings, latter 
particularly conspicuous on rostrum and chelae. 

Holotypic male, form I: Body subovate, depressed. Abdomen narrower 
than thorax ( 12.4 and 15.0 mm). Greatest width of carapace greater than 
depth at caudodorsal margin of cervical groove (15.0' and 10.1 mm). 
Areola moderately broad (4.9 times longer than wide) with number of 
deep punctations, three to six across narrowest portion. Cephalic section 
of carapace 1.7 times longer than areola (length of areola 36.6 per cent of 
entire length of carapace ) . Rostrum strongly acuminate, excavate dorsally 
with strongly convergent thickened margins devoid of marginal spines or 
tubercles; submarginal row of punctations deep basally becoming progres- 
sively shallower toward apex, median portion with a few scattered tuber- 
cles and distinct depression at level of caudal margin of orbit; acumen 
indistinctly delimited from remainder of rostrum and terminating in dor- 
sally directed tubercle; subrostral ridges strong and visible in dorsal aspect 
almost to base of acumen. Postorbital ridges heavy but short, grooved 
dorsolaterally, and terminating cephalically in low rounded tubercles. Sub- 
orbital angle obtuse. Branchiostegal spine very weak sinistrally and ob- 
solete dextrally. Carapace punctate dorsally, except in polished gastric 
area, and tuberculate laterally; tubercles especially strong in hepatic area 
and in row along ventral margin of cephalic portion of cervical groove; 
cervical spine lacking but several tubercles present in area usually support- 
ing it. Abdomen and carapace subequal in length. Cephalic section of tel- 



V ? OCT 3 1969 



284 Proceedings of the Biological Society of Washington 

son with two spines in each caudolateral corner, mesial spines movable. 
Dorsal surfaces of telson and uropods conspicuously setose. 

Epistome (Fig. 7) broader than long with subtriangular apex; lateral 
areas elevated, central area somewhat convex ventraUy and bearing many 
setae; caudomedian fovea present. Antennules of usual form with well 
developed spine on ventral surface of basal segment slightly distal to mid- 
length. Antenna broken but probably reaching fifth abdominal tergum. 
Antennal scale (Fig. 9) 2.3 times longer than broad, broadest at mid- 
length with widest lamellar area approximately 1.5 times width of 
thickened lateral portion, latter terminating in strong acute spine. 

Right chela (Fig. 12) somewhat depressed but with palm distinctly 
inflated; distal half of lateral margin of hand costate. All surfaces of palm 
and fingers punctate except opposable margin of latter and mesial margin 
of former. Mesial margin of palm with single row of seven depressed 
tubercles. Swollen ridge on dorsal surface of palm at base of dactyl very 
conspicuous. Ventral surface of palm with very large tubercle at mesial 
base of dactyl and smaller one proximolateral to it. Fingers gaping and 
both with well-defined longitudinal ridges dorsally and ventraUy. Op- 
posable margin of fixed finger with row of eight tubercles, proximal five 
much larger than distal three, and third from base largest; large tubercle 
present at lower level between sixth and seventh tubercles; one or two 
irregular rows of minute denticles extending distally from, base of fifth 
tubercle; corresponding margin of dactyl with nine tubercles, fourth from 
base largest and row of minute denticles extending distally from base of 
seventh tubercle. Mesial surface of dactyl entirely punctate. 

Carpus of right cheliped distinctly longer than broad with deep oblique 
longitudinal furrow dorsally; dorsal surface punctate; mesial surface with 
large spine slightly distal to midlength and smaller one proximal to 
midlength. Lower distal margin with large tubercle on lateral articular 
knob and massive spiniform tubercle mesial to it. 

Merus of right cheliped with mesial and lateral surfaces sparsely punc- 
tate, upper surface with one subdistal tubercle, ventrolateral margin with 
two tubercles, and ventromesial margin with row of seven, only six on 
sinistral chela. Row of three tubercles on ischium corresponding to mesial 
row on merus. 

Hooks on ischia of third pereiopods only (Fig. 8); hooks simple and 
not opposed by tubercle on basis but extending proximad of distal end of 
latter. Coxa of fourth pereiopod with prominent vertically disposed cau- 
domesial protuberance; fifth without prominences. 

Sternum deep between third, fourth, and fifth pereiopods and with 
prominent tuft of plumose setae between bases of third and fourth 
pereiopods. 

First pleopods (Figs. 2, 4, 6) symmetrical and reaching coxa of third 
pereiopods when abdomen is flexed ( See diagnosis for description ) . 

Morphotypic male, form II: Differs from holotype in following 
respects: mesial margin of palm with row of eight tubercles on dextral 
chela; opposable margin of fixed finger of chela with only five rounded 



New Georgia crayfish 285 

tubercles; corresponding margin of dactyl with row of eight; ventral sur- 
face of merus with mesial row of eight tubercles and ventrolateral margin 
with one spine on sinistral cheliped; ischium of sinistral cheliped with 
only two tubercles; hooks on ischia of third pereiopods and protuberances 
on coxae of fourth reduced in size; and first pleopod more shallowly em- 
bedded in sternum. ( See measurements. ) 

First pleopod ( Figs. 3, 5 ) with neither element corneous; mesial process 
fingerlike, rounded distally with only faint indication of bifid condition ex- 
isting in holotype, and directed caudolaterally and slightly distally ( in re- 
lation to shaft of appendage); central projection non-corneous, broadly 
rounded without any indication of subterminal notch, and directed 
caudally, its tip never extending caudally beyond tip of mesial process. 

Allotypic Female: Differs from holotype in following respects: mesial 
margin of palm of dextral chela with row of eight tubercles; opposable 
margin of fixed finger of sinistral chela with row of six tubercles; mesial 
surface of carpus with single major tubercle; ventrolatral margin of 
merus of dextral cheliped with row of three tubercles and ventromesial 
margin of sinistral cheliped with row of eight. Sternum between fourth 
pereiopods broad and moderately shallow. ( See measurements. ) First 
pleopods uniramus and barely reaching caudal margin of annulus ven- 
tralis. 

Annulus ventralis (Fig. 11) approximately 1.3 times broader than long, 
rather firmly fused to sternum cephalically, but caudal half slightly 
movable; lateral portions elevated (ventrally); cephalomesial areas with 
distinct median furrow extending caudally to midlength of annulus; sinus 
extending transversely toward sinistral wall, disappearing beneath it, 
there making U-shaped turn to cross median line, after which turning 
caudodextrally and then caudally to caudal margin of annulus. 

Type-locality: Sope Creek, tributary to the Chattahoochee River at 
Paper Mill Road 1.5 miles above mouth, Cobb County, Georgia. There 
the creek, some 70 feet wide and 2r-3 feet in depth, flows with a cascad- 
ing current over bed rock and scattered stones, the latter entrapping masses 
of filamentous algae. The steep shoreline is heavily wooded with such 
tree as Liriodendron tulipifera, Ulmus americana, Cornus flonda, and 
Acer rubrum. Associated with Cambarus howardi were Procambarus 
spiculijer (LeConte, 1856: 401) and Cambarus latimanus (LeConte, 
1856: 402). 

Disposition of Types: The holotypic male, form I, the allotypic female, 
and the morphotypic male, form II, are deposited in the Smithsonian In- 
stitution (nos. 129866, 129867, and 129868, respectively) as are the para- 
types, which consist of 6 ,J , form I; 6 ^ , form II; 11 $ ; 3 juvenile $ ; 1 
juvenile 2 ; and 1 2 with eggs. 

Size: The largest male, form I, has a carapace length of 31.8 mm, the 
largest female, 34 mm, and the smallest first form male, 27.6 mm. The 
former two specimens were collected from Sweetwater Creek and the 
latter from the type-locality. The largest specimen available is the allo- 
type. 



286 Proceedings of the Biological Society of Washington 

Color Notes: Cephalic portion of carapace and areola olive green dor- 
sally; margins of rostrum, postorbital ridges, and subrostral ridges scarlet. 
Branchiostegites greenish tan sometimes suffused with red. Terga of 
abdomen greenish tan margined caudally with narrow band of scarlet, 
pleura with cream borders; antennal scale margined in scarlet, otherwise 
green. Antennules and antennae scarlet, latter suffused with brown. Palm 
of chela light green with tubercles and swollen prominences scarlet, fin- 
gers green basally becoming progressively orange distally and terminating 
in scarlet tips. All legs greenish above fading to cream below but with 
scarlet markings at joints and scarlet dactyls. Sternum and bases of legs 
cream. 

Range and Crayfish Associates: This crayfish occurs in the upper pied- 
mont portion of the Chattahoochee Drainage System in Georgia where it 
is known to occur in only five localities : 

GEORGIA — Hall County, tributary of Balus Creek in east city limits of 
Oakwood ( 1 2 with eggs ) . Cobb County, type-locality, specimens listed 
above; Sope Creek at Barnes Mill Road SE of Marietta (1 $ II, 2 juv. $ , 
1 juv. $ ) ; Nickajack Creek at Camp Highland, 3.5 mi. S of Smyrna ( 2 $ 
I, 1 2 ). Douglas County, Sweetwater Creek at Factory Shoals Road 
south of Austell (1 $ I, 4 $ , 2 juvenile $ , 8 juvenile 2 ) ; Dog River at 
State Highway 5 SW of Douglasville (3 2 ) . In the Hall County locality, 
C. howardi was associated with C. latimanus, and in the remaining local- 
ities with both C. latimanus and Procambarus spicuUfer. 

Variations: The specimens from Douglas County differ from those of 
the type-locality in possessing, in general, areolae with more crowded and 
deeper punctations. This is especially noticeable in the specimens from 
Sweetwater Creek. Among those from the latter locality, the rostrum is 
less distinctly acuminate; the postorbital ridges terminate in spiniform 
tubercles; the lateral margin of the chela is costate throughout most of its 
length; and, like the larger male from Nickajack Creek, the fingers are 
distinctly longer and slenderer, and the palm much less inflated than in 
specimens from the type-locality. In general, the specimens from Dog 
River are more similar to those from Sweetwater Creek than to specimens 
from the type-locality. The single female from Hall County possesses a 
densely punctate areola, and, except for the chelae, which had been re- 
generated, resembles specimens from the type-locality. The small tubercle 
on the ventral surface of the palm of the chela, which is located proximo- 
lateral to the major tubercle in specimens from the type-locality, is situ- 
ated proximomesial to that tubercle in the specimens from Sweetwater 
Creek and Dog River. 

In specimens possessing a carapace length of at least 27.5 mm, the 
length of the areola constitutes 36.5 to 39.4 per cent of the carapace 
length, and in specimens having a carapace length of less than 27.5 mm, 
it comprises 34.3 to 35.9 per cent. The length of the areola ranges from 
3.0 to 4.9 times its width. 

Relationships: Cambarus howardi has its closest affinities with Cam- 
barus bartonii bartonii (Fabricius, 1798: 407), a highly variable species 



New Georgia crayfish 287 

which ranges from New Brunswick southward to northeast Georgia. It 
differs from the typical members of this species in possessing a more 
sharply acuminate rostrum, a somewhat densely punctate areola, which in 
larger specimens is usually somewhat longer, and in possessing a usually 
shorter, more robust chela. The most conspicuous difference occurs in 
the vivid scarlet markings on an olive to light green background. This 
crayfish, may eventually prove to be the southernmost subspecies of 
Cambarus bartonii. 

Life History Notes: First form males were collected in April, May, and 
October, and two females with eggs were taken in April and one in May. 

Etymology: In token of our appreciation for his interest in the water- 
ways of Georgia and the energy expended by him in preserving and im- 
proving them, we are pleased to name this species in honor of Mr. Ralph 
S. Howard, Jr., of the Georgia Water Quality Control Board. 

Remarks: This crayfish is an inhabitant of moderately to swiftly flow- 
ing streams and has been found only in areas where there is a rocky or 
rubble substrate. The type-locality is situated on the heavily polluted 
Sope Creek which takes its origin in the densely populated area im- 
mediately northeast of Marietta, Georgia. Almost at its source, it receives 
enrichment from storm sewers and an effluent from a sewage treatment 
plant. Six stations were established above the mouth of the creek ap- 
proximately (1) 8.0, (2) 7.3, (3) 5.8, (4) 5.0, (5) 3.0 and (6) 1.5 miles 
above it; just above station 5 it receives water from a relatively unpolluted 
tributary. On 21 July 1966, the percent saturation of oxygen at these 
stations was 87, 40, 23, 34, and 80.5, respectively. This crayfish, found 
only at stations 1 and 6, is believed to require an environment in which a 
high oxygen concentration exists, and, in view of these data, it seems pos- 
sible that were it not for the turbulent area immediately above Station 6 
which increases the dissolved oxygen concentration, Cambarus howardi 
would not be able to survive in the lower reaches of this stream. 

Cambarus ( Depressicambarus ) unestami new species 

Diagnosis: Body pigmented, eyes well-developed. Rostrum with con- 
vergent margins devoid of marginal spines or tubercles. Areola 3.5 to 4.7 
times longer than broad, comprising 32.6 to 37.4 percent of entire length 
of carapace, and bearing three to five punctations across narrowest part. 
Lateral surface of carapace lacking cervical spines or prominent tubercles. 
Suborbital angle prominent and subacute. Postorbital ridges short and 
rounded, with or without small cephalic tubercles. Antennal scale ap- 
proximately 2.2 times longer than broad. Chela with at least two rows of 
tubercles on mesial and mesiodorsal surfaces of palm, usually additional 
ones situated dorsolateral to these two rows; lateral margin of hand costate 
distally for at least half its length and both fingers with well-defined longi- 
tudinal ridge on upper surfaces. First pleopod (Figs. 14, 16, 18) of first 
form male with corneous central projection recurved at approximately 
125° to main shaft, not markedly tapering distally, with distinct subter- 
minal notch, and extending caudally beyond tip of mesial process; mesial 



288 Proceedings of the Biological Society of Washington 




Figs. 13-24. Camharus unestami new species (pubescence removed 
from all structures except for Fig. 14). 13, Lateral view of carapace of 
holotype. 14, Mesial view of first pleopod of paratypic male, form 1. 15, 
Mesial view of first pleopod of morphotype. 16, Caudal view of first pair 
of pleopods of holotype. 17, Lateral view of first pleopod of morphotype. 
18, Lateral view of first pleopod of paratypic male, form 1. 19, Epistome 
of holotype. 20, Bases of third, fourth, and fifth pereiopods of holotype. 
21, Antennal scale of holotype. 22, Dorsal view of carapace of holotype. 
23, Annulus ventralis and adjacent sternal elements of allotype. 24, Dorsal 
view af distal podomeres of chehped of holotype. 



New Georgia crayfish 289 

Table 2. Measurements (mm) of Cambarus unestami. 











Holotype 


Allotype 


Morphotype 


Carapace: 














Height 








10.8 


16.5 


10.7 


Width 








15.2 


21.2 


14.1 


Length 








29.6 


40.4 


27.8 


Areola: 














Width 








2.5 


3.7 


2.6 


Length 








10.5 


15.1 


9.5 


Rostrum: 














Width 








4.9 


6.0 


4.7 


Length 








5.7 


7.1 


6.6 


Chela: 














Length of 


inner 


margin 


of palm 


7.3 


10.9 


6.1 


Width of pahn 






10.2 


14.3 


8.6 


Length of 


outer 


margin 


of pahn 


22.0 


32.7 


19.4 


Length of dactyl 






13.2 


19.6 


11.3 



process inflated, directed caudolaterally and somewhat proximally and 
terminating in simple tip. Annulus ventralis (Fig. 23) somewhat asym- 
metrical with one lateral wall elevated and with ridge extending trans- 
versely from' opposite wall to disappear beneath elevated wall. Color 
brown to olive green with brownish black spots and bars, latter conspicu- 
ous on lateral surface of carapace and four broken longitudinal stripes on 
abdomen. 

Holotypic Male, Form I: Body subovate, depressed. Abdomen nar- 
rower than thorax (13.5 and 15.2 mm). Greatest width of carapace 
greater than depth at caudodorsal margin of cervical groove (15.2 and 
10.8 mm). Areola moderately broad (4.2 times longer than wide) with 
scattered punctations, four or five across narrowest portion. Cephalic 
section of carapace 1.8 times longer than areola (length of areola 35.5 per- 
cent of entire length of carapace ) . Rostrum acuminate, excavate dorsally, 
with convergent slightly thickened margins devoid of spines or tubercles. 
Submarginal row of punctations deep basally and becoming progres- 
sively shallower toward apex, median portion with few scattered tubercles 
and broad shallow depression at base; acumen not delimited from re- 
mainder of rostrum and terminating in small dorsally directed tubercle. 
Subrostral ridges moderately well-defined and visible basally, in dorsal 
aspect, to about midlength of rostrum. Postorbital ridges moderately 
strong, short, grooved dorsolaterally, and with cephalic extremity blunt; 
suborbital angle prominent and subacute. Branchiostegal spine reduced 
to small rounded tubercle. Carapace punctate dorsally and granulotuber- 
culate laterally; tubercles conspicuously strong in hepatic area and in row 



290 Proceedings of the Biological Society of Washington 

along ventral margin of cephalic portion of cervical groove. Cervical 
spines lacking, and area usually supporting them bearing tubercles little, 
if at all, larger than others on lateral surface. Abdomen longer than cara- 
pace (31.4 and 29.6 mm). Cephalic section of telson with two large and 
one small spine in caudodextral corner and two large ones in caudosinistral 
corner, medial of two larger spines on each side movable. Dorsal surfaces 
of uropod and telson strongly setose. 

Epistome (Fig. 19) broader than long, subtruncate cephalically, and 
lacking cephalomedian projection; central area slightly elevated ventrally, 
densely punctate but bearing few setae; caudomedian fovea almost ob- 
solete. Antennules of usual form with moderate spine on ventral surface 
of basal segment slightly distal to midlength. Antenna reaching sixth ab- 
dominal tergum. Antennal scale ( Fig. 21 ) 2.2 times longer than broad, 
broadest distal to midlength, widest lamellar area approximately 1.6 times 
broader than thickened lateral portion, later terminating in strong short 
subacute spine. 

Right chela ( Fig. 24 ) weakly depressed with inflated palm; distal three- 
fifths of lateral margin of hand costate. Mesial margin of palm with row 
of nine tubercles subtended dorsally by second row of four, three ad- 
ditional ones lateral to second row; remainder of dorsal surface of palm 
deeply pitted; ventral surface of palm mostly punctate with large sub- 
median distal tubercle; cluster of five smaller tubercles situated proximal 
to distal tubercle, single one at base of longitudinal ridge extending prox- 
imally from fixed finger, and another immediately ventrolateral to distal 
tubercle along mesial margin of palm. Fingers slightly gaping and both 
with well-defined ridges dorsally and ventrally. Opposable margin of fixed 
finger with row of five rounded corneous tubercles, fourth from, base in- 
jured but probably largest; large tubercle present on level below afore- 
mentioned row at base of distal two-fifths of finger; several rows of minute 
denticles extending from distal base of fifth tubercle to base of corneous 
tip of finger. Corresponding margin of dactyl with row of five 
rounded corneous tubercles, fourth from base largest; several rows of 
minute denticles extending from distal base of latter-mentioned tubercle 
to base of corneous tip of finger. Mesial surface of dactyl tuberculate and 
punctate along basal third, punctate distally. 

Carpus of right cheliped distinctly longer than broad with deep sinu- 
ous furrow dorsally; dorsomesial surface tuberculate; dorsolateral, lateral, 
and ventrolateral surfaces punctate; mesial surface with usual major spike- 
like tubercle, three smaller ones proximal to it, and one at its proximo- 
ventral base; distal mesioventral margin with prominent tubercle and 
group of three proximomesial to it; distal ventrolateral margin with prom- 
inent tubercle on condyle articulating with carpus and two small mar- 
ginal tubercles dorsolateral to it. 

Merus of right cheliped with mesial and lateral surfaces sparsely punc- 
tate, upper surface tuberculate, tubercles generally progressively larger 
distally; upper distal margin undulate; ventrolateral margins with row of 



New Georgia crayfish 291 

four tubercles, three of which spikehke, and ventromesial margin with 
twelve, eleven of which spikelike; oblique ventrodistal margin with two 
tubercles between terminal spikelike tubercles. Row of four tubercles on 
ischium corresponding to mesial row on merus. 

Hooks on ischia of third pereiopods only ( Fig. 20 ) ; hooks simple, not 
opposed by tubercle on basis, and extending proximad of distal end of 
latter. Coxa of fourth pereiopod with prominent vertically disposed 
caudomesial protuberance; fifth without prominences. Sinistral fifth leg 
absent, probably lost in early injury and not regenerated. 

Sternum deep between third, fourth, and fifth pereiopods and with 
prominent tufts of plumose setae between bases of third and fourth pereio- 
pods. ( See measurements. Table 2. ) 

First pleopods (Figs. 14, 16, 18) normally symmetrical and reaching 
coxae of third pereiopod when abdomen is flexed. (Actually, in the holo- 
type, the sinistral pleopod does not assume its characteristic attitude, prob- 
ably due to the absence of the corresponding fifth pereiopod. It is ab- 
normally twisted laterally and cannot be brought into its usual position 
between the bases of the caudal two pairs of pereiopods ) . ( See diagnosis 
for description. ) 

Morphotypic Male, Form II: Differs from holotype in following re- 
spects: cephalic extremity of postorbital ridges terminating in minute cor- 
neous tubercles; branchiostegal spine represented by small tubercle with 
acute corneous tip; lateral tubercles on carapace less prominent; cephalic 
section of telson with two spines in each caudolateral corner; epistome not 
truncate cephalically, but like holotype, lacking cephalomedian projec- 
tion; inner margin of palm of chela bearing row of seven tubercles, and 
irregular row of six immediately dorsolateral to it; tubercle at base of 
ventral longitudinal ridge of fixed finger lacking; ventral oblique ridge on 
palm at base of dactyl bearing prominent tubercle between major tubercle 
and articular socket; opposable margin of fixed finger with row of only 
four tubercles, third from base largest; row of minute denticles, broken by 
tubercles, extending distally from base of proximal tubercle; mesial surface 
of carpus of cheliped with only two tubercles proximal to spikelike tu- 
bercle; ventromesial row of tubercles on merus reduced to ten; hooks on 
ischia of third pereiopods and protuberances on coxae of fourth reduced in 
size. 

First pleopod distinctly more shallowly embedded in sternum than 
in holotype with tips of gonopod extending ventrally beyond ventralmost 
portion of coxae of fourth pereiopod; first pleopods (Figs. 15, 17) 
widely separated at base; mesial process heavy but tapering toward acute 
apex, disposed caudolaterally, and somewhat distally (in relation to the 
shaft of the appendage); central projection, non-corneous, broadly 
rounded, and directed caudoproximally, its tip extending caudally beyond 
tip of mesial process. 

Allotypic Female : Differs from holotype in following respects : cephalic 
section of telson with two spines in each caudolateral corner; right chela 



L 



292 Proceedings of the Biological Society of Washington 

regenerated, left chela with row of five tubercles above mesial row; three 
tubercles present ventrally opposite distal three tubercles in mesial row on 
palm; ventral surface of carpus with group of five tubercles proximal to 
marginal tubercles; merus with ventrolateral row of six tubercles and 
ventromesial row of thirteen. 

Sternum between fourth pereiopods broad and moderately shallow. 
First pleopods uniramus and reaching caudal margin of annulus. An- 
nulus ventralis (Fig. 23) approximately 1.5 times broader than long, 
firmly fused to sternum cephalically, but caudal half slightly movable; 
asymmetrical with high steeply sloping sinistral caudal wall; cephalo- 
mesial area with distinct median furrow extending caudally and curving 
dextrally into dextrally situated fossa; dextral wall high and curved; high 
ridge extending from sinistral margin transversely, continuing onto tongue, 
and latter disappearing beneath elevated dextral wall; sinus extending 
along caudal margin of tongue to median line and forming broad arc to 
end on caudal wall near median line. 

Type-locality: Daniel Creek, 2.5 miles west of Walker County line on 
State Route 143, Dade County, Georgia. There the creek, some 15 feet 
wide and 1.5 feet deep, flows with a moderate current over bed rock and 
a sandy bottom, the latter littered with stones and entrapped debris. The 
water is crystal clear and supports no macroscopic plant growth. The 
stream bed is located in a deep ravine in which Pinus sp. and Cornus 
florida are conspicuous elements. No other crayfish were found at this 
locality. 

Disposition of Types: The holotypic male, form I, the allotypic 
female with eggs, and the morphotypic male, form II, are deposited in 
the Smithsonian Institution (nos. 129863, 129864, 129865, respectively) 
as are the paratypes, which consist of 1 ^, form I; 14 $ , form II; 11 2; 
23 juvenile S ; 17 juvenile 2 ; and 1 2 with eggs. 

Size: The largest male, form I, has a carapace length of 31.4 mm, 
the largest female, the allotype, 40.2 mm, and the smallest and only other 
first form male available, the holotype, 29.6 mm. 

Color Notes: Ground color of carapace and abdomen brownish black 
dorsally fading ventrally to olive green and flecked with irregular brown- 
ish black splotches. Postorbital ridges brownish black. Lateral surface of 
carapace bearing brownish black "horn" (see Hobbs, 1958: 74) originat- 
ing in hepatic area and increasing in width caudally along branchiostegite 
at caudal margin of which almost reaching level of branchiocardiac 
groove. Terga of abdomen with dorsolateral linearly arranged series of 
oblique brownish black bars, similar series at bases of pleura. Telson and 
uropod with brownish black spots, Antennules and antennae dark brown. 
Chela brown with brownish black splotches on all podomeres distal to is- 
chium. Podomeres distal to ischium of remaining pereiopods olive green 
with brownish black spots. Basal podomeres of all pereiopods and sternum 
cream. 

Range and Crayfish Associates: This crayfish is known only from tribu- 
taries of the Tennessee River in Dade County, Georgia. While we can 



New Georgia crayfish 293 

cite only the type-locality, a tributary to Lookout Creek, as a definite 
locality for the species, we have juvenile specimens, which are tentatively 
assigned to this species, from a stream 0.7 miles east of the Alabama 
State line on Georgia State Highway 143 and from Rattlesnake Creek, 5.1 
miles northwest of U.S. 11 in Murphy Hollow. The only crayfish associate 
observed with this species was Camharus striatus Hay, 1902; 437, in the 
stream just east of the Alabama State line. 

Variations: The only available adult specimens of this species were 
collected in the type-locality and are remarkably uniform except for 
slight differences in the positions occupied by certain tubercles and an 
occasional loss or addition of a tubercle. Few differences extend beyond 
the range of variation occurring in the primary types. The number of 
tubercles along the mesial margin of the pakn of the chela varies from 
six to eight and the row immediately above it from four to six. The areola 
ranges from 32.6 to 37.4 percent of the total length of the carapace. 
Only the large allotypic female (carapace length, 40.4 mm) exhibits a 
percentage larger than 36.6; none of the other specimens have a carapace 
length greater than 32.9 mm. The length of the areola varies from 3.5 
to 4.7 times its width, but in the alltoype, it is only 4.1 times longer than 
broad. 

Relationships: Despite the fact that the chela of Camharus unestami 
is less depressed than that of any other member of the subgenus, the 
arrangement of the tubercles along the inner surface of the palm is 
typical of that of the other members; furthermore, the form of the 
terminal elements of the pleopod and the structure of the annulus 
ventralis indicate that its affinities are with the members of this segment 
of the genus. Unlike Camharus floridanus Hobbs, 1941: 114, C. latimanus 
(LeConte), C. reduncus Hobbs, 1956: 61, and C. striatus Hay, 1902: 437, 
this species possesses a subterminal notch on the distal end of the central 
projection of the first pleopod. It differs from C. halU Hobbs, 1968: 269, 
and C. obstipus Hall, 1959: 221, in lacking marginal tubercles on the 
rostrum. Finally, it differs from C. catagius Hobbs and Perkins, 1967: 
141, and C. sphenoides Hobbs, 1968: 262, in that the mesial process of 
the first pleopod does not extend so far caudally as does the central pro- 
jection. 

Life History Notes: A first form male and a female with eggs were 
collected in the type-locality on 5, May 1967, and a similar pair on 23, 
April 1968. 

Etymology: This species is named in honor of our mutual friend. Dr. 
Torgny Unestam, University of Uppsala, who was a most helpful and 
congenial companion to the senior author on a collecting trip in north 
Georgia when this species was discovered. 

Acknowledgments: We wish to express our appreciation to the follow- 
ing persons who have assisted in the collection of specimens of the two 
species described above: T. A. English, Jr., R. M. Gaddis, J. L. Ledbetter, 
J. L. Ledbetter, Jr., and Torgny Unestam. 



294 Proceedings of the Biological Society of Washington 

Literature Cited 

Fabricius, Johann C. 1798. Supplementum entomologiae systematicae. 
Hafniae, Proft. et Storch., pp. 1-572. 

Hall, Edward Taylor, Jr. 1959. A new crayfish of the genus Cambarus 
from Alabama (Decapoda, Astacidae). Journ. Tenn. Acad. 
Sci., 34(4): 221-225, 9 figs. 

Hay, William Perry. 1902. Observations on the crustacean fauna of 
Nickajack Cave, Tennessee, and vicinity. Proc. U.S. Nat. Mus., 
25(1292): 417-439, 8 figs. 

HoBBS, HoRTON H., Jr. 1941. Three new Florida crayfishes of the sub- 
genus Cambarus (Decapoda, Astacidae). Amer. Midi. Nat., 
26(1): 110-121, 2 pis. 

. 1956. A new crayfish of the genus Cambarus from North 

Carolina and South Carolina (Decapoda, Astacidae). Journ. 
Elisha Mitchell Sci. Soc, 72( 1 ) : 61-67, 11 figs. 

. 1958. The evolutionary history of the Pictus Group of the 

crayfish genus Procambarus. Quart. Jour. Fla. Acad. Sci., 21 
(1): 71-91, 20 figs. 

. 1968. Two new crayfishes of the genus Cambarus from Geor- 
gia, Kentucky, and Tennessee (Decapoda, Astacidae). Proc. 
Biol. Soc. Wash., 81: 261-274, 22 figs. 

, (in press). On the distribution and phylogeny of the cray- 
fish genus Cambarus. Virginia Polytechnic Institute Press. 

HoBBS, HoRTON H., Jr. AND Frank O. Perkins. 1967. A new burrowing 
crayfish from North Carolina (Decapoda, Astacidae). Proc. 
Biol. Soc. Wash., 80: 141-146, 11 figs. 

Leconte, John. 1856. Descriptions of new species of Astacus from 
Georgia. Proc. Acad. Nat. Sci., Phila., 7: 400-402. 



I 



! 



QH 

1 

B4X 



-322 3 October 1969 

PROCEEDINGS 



OF THE ( OCT 2 01969 ) 

BIOLOGICAL SOCIETY OF WASHINGTOHi^ ^^ 7/ 

A NEW SPECIES OF ILLEX FROM THE WESTERN 

ATLANTIC AND DISTRIBUTIONAL ASPECTS OF 

OTHER 7LLEX SPECIES (CEPHALOPODA: 

OEGOPSIDA).! 

By Clyde F. E. Roper,^ C. C. Lu,^ and Katharina Mangold^ 

Introduction: During the course of an examination of ceph- 
alopod specimens of the genus Illex from tropical waters of 
the western Atlantic, two of us ( KM and CFER ) discovered 
specimens that did not conform to the characteristics of previ- 
ously known species. While working at the Smithsonian Insti- 
tution on a large collection of Illex from the Chesapeake Bight 
region as part of a monographic study on the genus, one of us 
(CCL) encountered additional specimens of the undescribed 
species. 

Traditionally, two forms of Illex have been recognized in 
the North Atlantic; they have long, though not exclusively, been 
referred to as subspecies: Illex illecebrosus illecebrosus (Le- 
sueur, 1821 ) lq the western Atlantic from northern Brazil, the 
Caribbean, the Gulf of Mexico, north to Newfoundland and 
across northern European waters south to Bristol Channel; and 
Illex illecebrosus coindetii ( Verany, 1837 ) in the eastern Atlan- 
tic from the British Isles through the Mediterranean and south 
along the western African coast to Angola. The investigation 
that led to the discovery of the new species centered around an 
attempt to establish the taxonomic and distributional relation- 
ships of the presumed Illex illecebrosus illecebrosus in the trop- 
ical western Atlantic to the north Atlantic form of the spe- 
cies and to 7. illecebrosus coindetii in the eastern Atlantic and 



1 Contribution No. 28 from the Marine Sciences Research Laboratory, 

2 Division of Mollusks, Smithsonian Institution, Washington, D.C. 20560. 

3 Marine Sciences Research Laboratory, Memorial University, St. John's, New- 
foundland, Canada. 

* Laboratoire Arago, Banyuls-sur-Mer, France. 

22— Proc. Biol. Soc. Wash., Vol. 82, 1968 (295) 



296 Proceedings of the Biological Society of Washington 



* II lex oxygonius 
®Holotype 

• Illex coindetii 




Figure 1. Capture-localities of Illex species in the middle western 
North Atlantic. 



the Mediterranean. The result of this investigation are reported 
here: the subspecies of Illex, I. illecebrosus illecebrosus, I. il- 
lecehrosus coindetii, and the recently estabUshed I. illecebro- 
sus argentinus (Castellanos, 1960) are demonstrated to have 
status as full species; the new species is described and com- 
pared with the other species of Illex; I. coindetii is recorded for 
the first time from the western Atlantic in warm waters; and 
general distributional ranges are discussed. A full and detailed 
revision of the systematic and zoogeography of all species of 
Illex is now in progress ( CCL ) . 

Acknowledgments: The co-authors wish to acknowledge the 
support of the following individuals and institutions: Speci- 
mens from the Chesapeake Bight region have been provided by 



The genus Illex 297 

by J. Davis and J. Musik from the Virginia Institute of Marine 
Science; from the Gulf of Mexico and the Gulf of Guinea by 
the U. S. Bureau of Commercial Fisheries through H. BuUis; 
from the Gulf of Mexico and Florida waters by G. L. Voss, In- 
stitute of Marine Sciences, University of Miami. The remain- 
ing study material came from the collections of the Smithsonian 
Institution and from the authors' collections. Funds have been 
provided through grants from the Centre National de la Re- 
cherche Scientifique (KM ), the National Research Council and 
the Fisheries Research Board of Canada to F. A. Aldrich 
(CCL), and the Smithsonian Research Foundation (CFER). 

G. L. Voss kindly provided working space and specimens 
during a visit to the Institute of Marine Sciences by one of us 
(KM). F. A. Aldrich kindly provided continued support for 
and interest in the studies on the systematics of Illex (CCL). 
The manuscript had been read and valuable comments pro- 
vided by R. E. Young, J. Rosewater, G. L. Voss and F. A. 
Aldrich. The illustrations of the new species have been ren- 
dered by J. Schroeder. The chart of distribution was prepared 
by M. Sweeney. We gratefully acknowledge the assistance of 
these individuals. 

Measurements and Abbreviations: Most of the measurements 
(in mm) and indices used in this paper are those commonly 
used for squid, with dorsal mantle length (ML) the standard 
against which other measurable characters are compared. Ab- 
breviations of most measurements and indices are presented in 
Voss ( 1963 ) . Some additional measurements have been taken 
to aid in the description of the species and to help define sex- 
ual dimorphism. Normally, mantle width ( M W ) is given as a 
single value that represents the greatest width of the mantle; in 
this study three values for mantle width are utilized: 

MWi — width at mantle opening 

MW2 — width at point half-way between anterior insertion of 

fins and anterior end of mantle ( dorsal side ) 
MW3 — width at point in line with anterior insertion of fins 
HL — length of head along dorsal midline from posterior 

ridge to V-shaped juncture between arms I. 

The fin angle is the angle that the straight posterior border of 






298 Proceedings of the Biological Society of Washington 

Table 1. Measurements (in mm) of Illex oxygonius new species 





1 

1 


C/3 

> 


g 
> 


1 
> 


X 
fe 




C/3 
> 


1 

o 


> 


2 
> 


^ 
^ 


Sta. No. 


6801 


167 


167 


167 7296 7281 


167 6801 


167 


167 


167 


Sex 


$ 


$ 


$ 


$ 


$ 


$ 


2 


2 


2 


2 


9 


DML 


135 


160 


176 


178 


205 


207 


174 


181 


184 


196 


210 


VML 


120 


144 


172 


166 


192 


200 


174 


174 


176 


189 


210 


MWi 


26 


27 


32 


34 


39 


39 


31 


29 


33 


33 


37 


MWa 


22 


22 


23 


30 


32 


31 


30 


28 


29 


30 


39 


MWs 


15 


13 


19 


18 


19 


19 


22 


17 


18 


21 


30 


FL 


60 


69 


74 


80 


92 


94 


79 


77 


83 


84 


95 


FW 


70 


70 


84 


90 


99 


98 


99 


78 


91 


99 


107 


Fin angle 


35° 


30° 


33° 


31° 


27° 


28° 


38° 


27° 


34° 


34° 


30° 


HL 


25 


27 


29 


35 


33 


31 


28 


26 


29 


26 


39 


HW 


27 


33 


33 


36 


47 


44 


33 


29 


32 


33 


41 


Ai 


56 


58 


62 


74 


87 


88 


65 


51 


58 


57 


74 


A„ 


68 


72 


86 


96 


109 


108 


78 


66 


71 


68 


94 


Am 


68 


70 


88 


97 


103 


108 


74 


66 


70 


72 


98 


Aiv (normal) 


62 


63 


79 


80 


92 


88 


70 


58 


65 


64 


83 


Aiv (hect.) 


75 


65 


85 


90 


103 


103 












Length of 
























hect. portion 22 


20 


26 


27 


32 


33 












TL 


115 


115 





112 


120 


125 


108 


121 


112 


104 


135 


CL 


54 


54 


— 


51 


53 


61 


47 


58 


50 


47 


61 


Si 


1.9 


2.1 


2.7 


3.1 


3.0 


3.0 


2.3 


2.0 


2.3 


2.2 


2.8 


Sii 


3.5 


3.6 


4.0 


4.5 


5.2 


5.0 


3.2 


3.5 


3.2 


3.1 


4.3 


Siii 


3.5 


3.5 


4.0 


4.5 


5.0 


5.0 


3.2 


3.4 


3.0 


3.1 


4.1 


Siv 


1.9 


1.9 


2.5 


2.7 


2.8 


2.7 


2.1 


1.9 


2.2 


2.0 


2.6 


Siv (hect.) 


1.7 


1.9 


2.3 


2.7 


2.8 


2.4 












Sciu. 


2.3 


2.5 




3.4 


4.0 


4.0 


2.9 




2.7 


2.9 


4.0 


Hect. arm 


L 


L 


L 


L 


R 


L 












Nid. gl. 














96 


76 


23 


— 


106 



one fin, exclusive of the lateral lobe, makes with the longitu- 
dinal axis (mid-dorsal line) of the body. 

Table 1 lists the measurements of Illex oxygonius, and Table 
2 presents the ranges and means of the indices. 

Several abbreviations of ships and institutions are used in ref- 
erence to specimens : 
FH Fish Hawk 
G Grampus 
O Oregon 
VIMS Virginia Institute of Marine Science 



The genus Illex 299 

Family OMMASTREPHIDAE Steenstmp, 1857 

Genus Illex Steenstmp, 1880 

Type species: Illex illecehrosus (Lesueur, 1821). 

Illex oxygonius, new species 

Plates 1-5, lOA 

LIST OF MATERIAL 



Sex 


ML, 
mm 


Station 


Location 


Date 




Depth, 
m 


Holotype: 












$ 


207 


FH 7281 


24°13'N81°58'W 


14 II 


1902 


555 


Paratypes: 












$ 


205 


FH 7296 


24°22'N81°48'W 


26 II 


1902 


222 


$ 


160 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


$ 


135 


O 6801 


29°39.5'N 80°08.5'W 


20 VII 1967 


383 


2 


181 


O 6801 


29°39.5'N 80°08.5'W 


20 VII 


1967 


383 


S 


176 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


9 


210 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


Other material: 










$ 


228 


G-line 2 


38°52'N 72°58'W 


8 X 


1892 


155 


$ 


178 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


5 


174 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


? 


184 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


5 


196 


VIMS 167 


38°27'N 73°23'W 


25 VIII 


1967 


190 


$ 


170 


VIMS 188 


36°18'N 74°52'W 


31 VIII 


1967 


68 


2 


181 


VIMS 177 


37°22'N 74°47'W 


27 VIII 


1967 


50 


$ 


164 


VIMS 166 


38°34'N 73°33'W 


25 VIII 


1967 


73 


2 


136 


O 1011 


24°28'N 83°25'W 


14 IV 


1954 


364 



Diagnosis: Fin angle acute (25°-35°); fin width equal to fin length; 
mantle narrow, drawn out posteriorly; dorsal mantle lobe conspicuous, 
pointed in males; hectocotylized portion of arm IV long, 3 papillae on 
dorsal row; cone at oral end of cement body of spermatophore funnel- 
shaped. 

Description: Mantle long, slender, broadest anteriorly, drawn out 
posteriorly into long attenuate tip (pi. lA). Mantle width decreases 
markedly posteriorly in males, less so in females especially at maturity 
( Table 2 ) . Ventro-lateral lobes on mantle opening inconspicous, rounded; 
dorsal lobe conspicuous, pointed ( pi. IB ) . Thin, membranous flange en- 
circles mantle opening. Mantle wall thin for an ommastrephid. Fins 
narrow, drawn out to acute point posteriorly; width (48-50 percent of 
ML) about equal to length (44 percent of ML); ratio of length to width 
= 1:1.01 to 1:1.25; slightly concave posterior borders form acute angle of 
25°-35° (average about 30° in males, 32° in females); lateral borders 
rounded; anterior lobes well developed (pi. IB). 



300 Proceedings of the Biological Society of Washington 



Table 2. 



Ranges and means of indices of Illex oxygonius new species 
(n = ll; 6 males, 5 females) 





Males 


Females 




Mean 


Range 


Mean 


Range 


MWiI 


18.3 


17-19 


17.4 


16-18 


MW2I 


15.2 


13-17 


16.4 


15-19 


MW3I 


9.7 


8-11 


11.4 


9-14 


HLI 


17.1 


15-20 


15.6 


13-19 


HWI 


20.7 


19-23 


17.8 


16-20 


FLI 


44.0 


42-45 


44.2 


43^5 


FWI 


48.2 


44-52 


50.4 


43-57 


FL/FW 


1/1.10 


1/1.01-1/1.17 


1/1.13 


1/1.01-1/1.25 


Fin angle 


30.7 


27-35 


32.6 


27-38 


All 


39.8 


35^3 


32.2 


29^37 


An I 


50.5 


45-54 


40.0 


35-45 


Ami 


50.2 


44-55 


40.2 


3&-47 


Aivl 


43.9 


39^6 


36.0 


32^0 


Aiv(hect.)I 


49.3 


41-56 






Hect. portion 










percent of arm 


30.8 


29-33 






Si I 


1.49 


1.31-1.74 


1.20 


1.10-1.33 


Siil 


2.41 


2.25-2.59 


1.83 


1.58-2.05 


Sml 


2.39 


2.19-2.59 


1.77 


1.58-1.95 


Sivl 


1.37 


1.19-1.52 


1.14 


1.02-1.24 


Snect. I 


1.30 


1.16-1.52 






Sciubl 


1.81 


1.56-1.95 


1.63 


1.47-1.90 



Funnel well developed, set in deep funnel groove; lateral adductor 
muscles conspicuous, strong, rodlike (pi. lA); anterior adductors thin, 
broad, sheetlike. Dorsal funnel organ large (pi. 3A); posterior limbs 
fleshy, thickened, broad anteriorly, tapered, bluntly pointed posteriorly; 
lateral shoulders conspicuous; anterolateral borders concave; apical papilla 
spear-head shaped. Sharp, thin ridges extend from anterior mid-portion 
of limbs nearly to apex. Ventral pads oblong; lateral borders strongly 
curved; medial borders nearly straight. Fuimel valve broad. 

Funnel locking-cartilage large, strong, inverted T-shaped (pi. 2B). 
Medial, longitudinal groove narrow, relatively shallow; groove deepens 
and widens posteriorly to form deep, broad pit. Strongly developed 
cartilaginous knobs converge, nearly meet posterior to deep pit; lateral 
knob rounded, medial knob more pointed with narrow ridge extending 
dorsally into pit; pit undercuts anterior walls of knobs. Groove between 
knobs narrow, relatively deep, opens into lateral groove. Lateral groove 
very broadly inverted V-shaped, deepest anteriorly against posterior walls 
of knobs, shallows along posterior margin of locking-cartilage. Entire 
cartilage bordered by thin, muscular flange. Mantle locking-cartilage 
( pi. 2C ) strong, compliment of funnel lock. Longitudinal ridge low, nar- 



The genus lUex 



301 



)4 1 1 




Plate 1. I//ex oxygonius new species, Holotype, FH 7281, male, 207 
mm ML. A. Ventral. B. Dorsal. 



row anteriorly, expands to large, sharply defined, swollen bulb posteriorly; 
posterior wall very deeply undercut toward base on mantle. Thin ridge 
extends from bulb to lateral ridge. Lateral ridge narrow, open V-shaped; 
anterior walls drop sharply into open grooves, posterior walls taper to 
mantle wall. 



302 Proceedings of the Biological Society of Washington 






Plate 2. Ulex oxygonius new species, Holotype. A. Left tentacular 
club. B, C. Funnel and mantle components of locking cartilage. D. Nuchal 
locking cartilage. E. Right arm III. 



Head large, broad, width about 20 percent of ML in males, 17 percent 
in females; length about 17 percent in males, 15 percent in females. Three 
flaplike nuchal folds present; nuchal cartilage long, broad and roimded 
anteriorly, narrow, tapered posteriorly, terminally embedded in muscle; 



The genus Illex 303 

central groove deep, lateral ridges distinct (pi. 2D). Eye openings sub- 
circular, dorsoventrally elongate; anterior sinus deep, conspicuous. 

Arms long, attenuate; order 2 ^ 3 > 4 > 1; longer in males than in 
females, especially II and III ( Table 1 ) ; arms II, III robust; IV less robust; 
I least robust, very slender ( pi. 1 A, B ) . Swimming keels very low, weak 
on distal half of I; higher, better developed, full length of II; best devel- 
oped on III, broadest in proximal quarter; low, second best developed, 
full-length of IV. Protective membranes well-developed, particulary ven- 
tral membranes; trabeculae long, very strong, arise from base of sucker 
stalks, form high points along membranes. Suckers of arms larger in 
males than in females, especially on II and III (Table 2). Suckers small 
on I and III; about 6 pairs enlarged on II and III of male ( pi. 2E ) . 

Arm sucker dentition. Largest rings of Arm I ( pi. 4A ) with low, broad 
plate on proximal 1/3, 3—5 low, truncate or slightly rounded teeth later- 
ally, becoming narrower, longer distally; distal median tooth enlarged, 
elongate, more pointed; teeth, especially median tooth, more truncate, 
blunt on basal sucker pairs, more pointed on distal pairs. Largest rings 
of II ( pi. 4B ) with 2 low, broad, flat teeth proximally, 5-7 teeth laterally, 
broader, more truncate proximally, narrower, more rounded distaUy; 
distal median tooth greatly enlarged, triangular, pointed. Smaller rings 
smooth proximally and with fewer teeth, 5-6 long, narrow teeth distally, 
grade in size to median tooth. Ring dentition of III about as II; broad, 
low single or bifurcate plate proximally, 5-7 lateral teeth graded distally, 
median tooth very enlarged, elongate, pointed (pi. 4C). Smaller rings 
with broad, low plates proximally, 1-2 roimded teeth laterally, 2-3 long 
pointed teeth laterally graded to longest median tooth. Largest rings of 
IV with broad, low proximal plate, 3-5 rounded to truncate lateral teeth 
(may be longer, narrower distally), elongate bluntly pointed or rounded 
distal median tooth. 

Left or right ventral arm hectocotylized (pi. 3B), longer (average 
about 6 percent ) , more robust than non-hectocotylized arm IV; modified 
tip 29^3 percent ( average 31 percent ) of arm length; distal suckers re- 
duced in size, rows separated. Suckers of dorsal row give way to 3^ conical 
papillae; remnant of protective membrane disappears distal to third 
papilla; next distal 1-2 protuberances become flattened, followed by series 
of transversely broad, thin, nearly truncate flaps; flaps gradually reduced 
in size distally, finally giving way at arm tip to series of minute papillae 
(28 recognizable flaps on holotype-exclusive of proximal 3 papillae and 
9 distal precursors ) . Suckers of ventral row give way to conical, nipple- 
like papillae that extend, gradually reduced in size, to tip of arm (42 
papillae in holotype). Low, weak zigzag ridge between papillae and 
flaps. Aboral keel broadly expanded along modified portion of hectocoty- 
lus tip. Trabeculae not modified as fringed lobes. 

Tentacles robust, relatively short; stalks naked. Clubs expanded, long, 
about 28-30 percent of ML (pi. 2A). Distinct carpal cluster absent; 
suckers in carpal area small, biserial; carpal knobs lacking. Manal suckers 
tetraserial; suckers on lateral rows small; medial sucker rows arise distal 



304 Proceedings of the Biological Society of Washington 





The genus Illex 305 

to the proximal 5 carpal suckers; proximal 3-5 medial suckers small, 
gradually increasing in diameter; about 15 medial suckers on manus greatly 
enlarged; maximum size greater in males than in females; manus termin- 
ates abruptly. Dactylus distinct, slender; suckers in 8 or 9' rows, extremely 
numerous, minute; suckers near tip slightly enlarged; tip with narrow, 
suckerless flange. Swimming keel along entire tentacular stalk aborally, 
troadest proximal to club, diminishes along carpal and manal region, ex- 
pands to broad keel along dactylus. Lateral angles distinct on oral surface 
of stalk and continue as broad heavily supported protective membranes 
along club, diminish significantly along dactylus. Dual V-shaped trabec- 
ulae arise from enlarged common base with lateral suckers along manus. 

Club sucker dentition. Sucker rings of carpal area with low, broad plate 
on proximal half, 4-6 small teeth on distal half; lateral teeth broad, 
rounded; distal teeth narrow, bluntly pointed. Proximal-most lateral manal 
suckers with proximal plate, 1-2 low, rounded lateral teeth, 4-7 long, nar- 
row pointed distal teeth; distal lateral suckers toothed around entire 
margin with 5-8 low, rounded proximal teeth, 12^14 longer, sharply 
pointed lateral and distal teeth, occasional small pointed secondary tooth 
between primary teeth. Proximal medial manal suckers (first 5-7) with 
low proximal plate, 1-2 low, rounded, lateral teeth, 1-2 slightly narrower 
distolateral teeth, 1 buntly pointed median tooth; next distal 4-5 enlarged 
sucker rings with 7-13 teeth around margin — low, flat proximally, narrow, 
rounded distally, no enlarged or pointed medial tooth ( pi. 4E ) . Dentition 
lacking, rings entire on 7-8 distal enlarged manal suckers. Proximal dactyl 
sucker rings with 4-6 long pointed distal teeth; teeth become more blunt, 
rounded on rings of middle dactylus suckers; teeth on suckers at distal 
tip few, minute. 

Gladius ( pi. 5A ) long, slender, bluntly pointed anteriorly; median ridge 
and heavy lateral rods extend length of rhachis, converge posteriorly, fuse 
near vane, continue to tip. Vane short, narrow; forms short, hollow conus; 
only extreme tip solid. 

Connectives of buccal membrane attach to dorsal, dorsal, ventral, dorsal 
borders of arms I-IV respectively. 

Beaks ( pi. 4G, H ) . Rostrum of upper beak strong, sharp, long, heavily 
pigmented; dorsal hood very weak, thin, short, Hghtly pigmented; notch 
and slight ridge separate rostrum and rostral lamella; cutting edge of rostral 
lamella straight or slightly curved, smooth; palatine lamella large, long, 
deep, pigmented anteriorly, dorsal crest only slightly curved. Rostrum of 
lower beak relatively long, pointed, heavily pigmented, inner edge curved; 
rostral lamellae narrow, short, wing lobate, thin; gular lamellae large, 
crest strong, posterior corner long, pointed, curved; rostral width broad. 



Plate 3. Illex oxygonius new species. A. Open funnel showing funnel 
organ and funnel valve. Holotype. B. Hectocotylus, right arm IV. Para- 
type, FH 7296, male, 205 mm ML. 



306 Proceedings of the Biological Society of Washington 




^^ ci d^ ^ W ^ 





Plate 4. lllex oxygonius new species. A-E. Sucker rings from arms 
I-IV and manus of club, respectively. Holotype. F. Radula, G. Upper 
beak, H. Lower beak. Paratype, FH 7296. 



Radula (pi. 4F) with 7 transverse rows of teeth; rhachidian with long 
central tooth, blunt lateral cusps; first lateral tooth with blunt lateral 
cusp; second lateral tooth curved, no lateral cusp; third lateral tooth long, 
curved, scythe-shaped; marginal plaques lacking. 

Spermatophore (pi. 5B-D, pi. lOA) with sperm mass of about 60 per- 
cent of total length, cement body about 15 percent of total length. Cone at 



The genus Illex 



307 




Plate 5. Illex oxygonius new species. Paratype, VIMS 167, males, 176 
mm ML. A. Gladius. B-D. Spermatophore. B. Oral end. C. Mid-section; 
cement body ejaculatory apparatus. D. Entire spermatophore. 



308 Proceedings of the Biological Society of Washington 




Plate 6. Illex illecebrosus (Lesueur, 1821). A. Male, 252 mm ML. 
B. Female, 268 mm ML. Newfoundland (from Mangold, et al., in press). 




309 



Plate 7. Illex coindetii (Verany, 1837) A. Male, 166 mm ML. B. Fe- 
male, 95 mm ML. Western Mediterranean (from Mangold, et al., in 
press ) . 



310 Proceedings of the Biological Society of Washington 





Plate 8. Illex argentinus ( Castellanos, 1960). A. Female, 187 mm 
ML. B. Male, 172 mm ML Argentina. 



oral end of cement body funnel-shaped, connected aborally by short neck; 
oral tube relatively wide. 

Color (in isopropyl alcohol). Reddish-brown chromatophores cover 
entire body, more widely spaced ventrally, thickly packed dorsaUy; deep 
purple-red stripe along dorsal mid-line of mantle. 



The genus Illex 



^^^i^feSKtiJ^^j^ 






311 



B 



?^^4*X!ii#'%S^ 



fe*^.^ 




Plate. 9. Hectocotyli of Illex species. A. I. illecehrosus, 252 mm ML; 
Newfoundland. B. I. argentintis, 167 mm ML; Argentina. C. I. coindetii, 
166 mm ML; western Mediterranean. D. I. coindetii, enlargement of a 
portion of C. showing fringed trabeculae. 

Holotype: United States National Museimi (577000). 

Type locality. 24°13'N 81°58'W in Florida Current about 22 nautical 
miles SSW of Key West, Florida; Fish Hawk station 7281, 14 February 
1902, 555 m. 

Distribution: Western Atlantic: off Middle Atlantic States, Florida 
Current, Gulf of Mexico (fig. 1). Taken in bottom trawls at depths of 
50-555 m. 



312 Proceedings of the Biological Society of Washington 

f il 








The genus Illex 313 

Etymology: The name of the species is derived from the Greek word 
"oxygonios" meaning acute angled. This term refers to the acute angle of 
the fins in I. oxygonius. 

Discussion: Illex oxygonius. Illex oxygonius exhibits a considerable 
sexual dimorphism in several characters (Table 2). The width of the 
mantle in males decreases sharply posteriorly, while that of females de- 
creases gradually; this gives females a fuller, more robust appearance along 
the middle and posterior portions of the mantle. Head size differs be- 
tween sexes; males have longer, wider heads than females. Fin width in 
females tends to be slightly wider than in males; this results in a slightly 
greater fin angle. All arms in males are longer than the corresponding 
arms in females; arms II and III are especially more elongate and more 
robust. The maximum size of suckers on all arms, particularly on arms II 
and III, is greater in males than in females. Males have larger tentacular 
club suckers than do females. 

While few specimens are currently available, it appears that J. oxygon- 
ius generally may inhabit depths greater than around 100-200 m, partic- 
ularly in lower latitudes where it is recorded from 222 to 555 m. 

All males of I. oxygonius available are mature with completely devel- 
oped hectocotyli and Needham's sacs filled with completely formed sper- 
matophores; males range from 135-228 mm in ML. At least some females 
become fully mature by 174 mm ML with swollen nidamental glands of 
96 mm length and completely filled ovaries. 

Comparison of species. Steenstrup (1880) erected the genus Illex to 
encompass two distinct species, I. illecebrosus (Lesueur, 1821) and I. 
coindetii (Verany, 1837). Pfeffer (1912) had access to a large collection 
of Illex from a number of localities in the Atlantic; he formed the opinion 
that illecebrosus and coindetii were subspecies. Pfeffer's subspecific 
designations have been followed since (e.g., Degner, 1925; Grimpe, 1921, 
1924, 1925; Adam, 1939, 1952). Acceptance of the subspecies concept, 
however, has not been universal (Naef, 1916, 1921, 1923; Voss, 1956; 
Clarke, 1966). 

The taxonomic problems in the genus Illex have been further com- 
plicated by the absence of holotypes. In fact, no type specimen was kept 
for illecebrosus by Lesueur (1821), and the type of coindetii is no longer 
extant (Voss, 1962). Castellanos (1960) described a new species called 
Ommastrephes argentinus which was later placed in Illex as the third sub- 
species oi illecebrosus (Castellanos, 1964). 

The comparisons of species are based on a large sample of specimens of 
Illex from various localities. Specimens of illecebrosus have been ex- 
amined and confirmed as conspecific from Newfoundland, and Labrador, 
Gulf of Maine, Middle Atlantic States, and off north Florida. Specimens 



Plate 10. Spermatophores of Illex species. A. I. oxygonius. B. I. il- 
lecebrosus. C. 7. coindetii. D. 7. argentinus. 



314 Proceedings of the Biological Society of Washington 

of coindetii come from the Mediterranean, Gulf of Guinea, West Africa, 
Gulf of Mexico and the Caribbean; we have examined and compared the 
specimens from these areas and have concluded that they are conspecific. 

Illex oxygonius may be distinguished from other species of Illex by a 
number of characters. Information on distinguishing characteristics has 
been accumulated from a large sample of specimens of illecehrosus and 
coindetii from the North Atlantic; information on argentius from South 
American waters, however, is available from only eleven specimens of 
which data from eight were presented in Mangold, et al. ( in press ) . Sev- 
eral of the characters are presented as indices of morphometric features; the 
mean values are given followed by the ranges in parentheses. In general, 
the indices tabulated are of illecehrosus from Newfoundland and of coin- 
detii from the western Mediterranean ( Mangold, et al,, in press), but other 
characters used include, in addition, illecehrosus from the Chesapeake 
Bight region and coindetii from West Africa, the Gulf of Mexico and 
Caribbean. Males and females of illecehrosUrS, coindetii and argentinus are 
illustrated in plates 6-8 respectively. 

Head width. The head width of oxygonius in general is greater than 
that of other species, with the exception of the male of coindetii. The 
males of oxygonius have an average head width index of 20.7 ( 19^-23), of 
illecehrosus 15.2 (14.5-17.1), of coindetii 20.0' (17-22.4), and oi urgen- 
tinus 15.9 ( 13.0^18.6); the values for females are, respectively: 17.8 ( 16- 
20); 14.9 (13.8-16.6); 15.5 (15.0-16.2); 14.3 (12.4-16a). Sexual di- 
morphism in head width occurs in all species except illecehrosus. 

Arms. Males and females of oxygonius have the smallest indices of all 
arms of the four species : 

I. oxygonius I. illecehrosus I. coindetii I. argentinus 

Male 

Arm I 39.8(35-43) 42.8(42.5-42.8) 54.9(42.2-63.8) 59.2(57.1-63.0) 

II 50.5(45-54) 51.4(51.8-55.3) 71.9(55.1-87.7) 73.0(68.7-77.8) 

III 50.2(44-55) 52.3(50.4-50.6) 71.4(53.3-87.7) 72.4(69.4-75.9) 

IV^ 43.9(39-46) 45.3(43.9-46.2) 52.4(48.2-77.6) 63.0(60.1-67.3) 

Female 

Arm I 32.2(29-37) 38.1(38.3-44.2) 43.8(39.7-43.7) 44.1(39.4-47.7) 

II 40 (35-45) 45.3(44.1-55.2) 56.3(56.2-56.5) 53.1(48.2-57.0) 

III 40.2(36-47) 46.2(44.1-53.3) 56.1(55.3-56.6) 53.2(47.6^56.8) 

IV 36 (32-40) 41.1(41.5-47.3) 50 (49.2-51.0) 47.1(41.8-50.0) 

1 Data for normal arm, not hectocotylus. 

The figures show that the males have longer arms than the females, 
particularly in oxygonius, condetii and argentinus; the sexual dimorphism in 
arm lengths of illecehrosus is less pronounced. Arms II and III of male 
oxygonius, coindetii and argentinus are more robust than corresponding 
arms in females; this is especially so in coindetii. Arms II and III of male 
illecehrosus are not enlarged. The largest suckers on arms II and III of male 



The genus Illex 



315 



oxygonius, coindetii and argentinus are conspicuously larger than those in 
females; no such enlargement occurs in illecehrosus: 





/. oxygonius 


7. coindetii 


I. argentinus 


I. illecehrosus 


Male 










Arm I 


1.49(1.31-1.74) 


1.74(1.28-2.22) 


1.64(1.58-1.72) 


1.17(0.93-1.54) 


II 


2.41 (2.25-2.59) 


3.44 (2.26-4.3^) 


3.21 (2.99-3.33) 


1.61(1.52-1.74) 


III 


2.39(2.19-2.59) 


3.29(2.15-3.99) 


2.97(2.88-3.07) 


1.62(1.48-1.79) 


IVi 


1.37(1.19-1.52) 


1.52(1.08-2.02) 


1.53(1.50-1.59) 


0.98(0.91-1.12) 


Female 










Arm I 


1.20(1.10-1.33) 


1.32(1.08-1.62) 


1.28(1.06-1.45) 


1.14(1.04-1.23) 


II 


1.83(1.58-2.05) 


2.06(1.78-2.54) 


2.01(1.59-2.18) 


1.64(1.37-1.82) 


III 


1.77(1.58-1.95) 


2.00(1.67-2.45) 


1.98(1.53-2.18) 


1.54(1.31-1.72) 


IV 


1.14(1.02-1.24) 


1.15(1.00-1.27) 


1.15(0.89-1.32) 


1.06(0.90-1.29) 



^ Data for normal arm, not hectocotylus. 

Beaks. The beaks of the three species differ in a number of features: 



Feature 


7. oxygonius 


7. illecehrosus 


7. coindetii 


Upper beak: 








Rostral hood 


short, very thin 
and weak 


long, strong 


long, strong 


Outline of rostral 


smooth, straight or 


serrated 


smooth 


lamella 


slightly curved 






Jaw angle 


small 


large, with tooth 


small 


Rostrum 


short 


long 


long 


Palatine lamella 


long, deep; crest 


short, shallow; 


short, shallow; 




straight 


crest curved 


crest curved 


Rostral lamella 


long 


short 


short 


Lower beak: 








Cutting edge of 


curved, long 


straight, short 


straight, long 


rostrum 








Rostral lamella 


short, narrow. 


long, wide, no 


long, wide, no 




lobate; irregular 


lobe; regular 


lobe; slightly 




outline 


outline 


irregular outline 


Gular lamella 


long, pointed 


short, blunt 


short, blunt 


Rostral width 


wide 


narrow 


narrow 



Features of the mantle. In Illex oxygonius and coindetii the widest 
part of the mantle is at the mantle opening except in fully mature fe- 
males because of the great enlargement of the nidamental glands and 
ovary. The mantle in oxygonius is long and narrow and tapers evenly to 
a point posteriorly, while that of condetii is shorter, proportionally less 
narrow, and less drawn out posteriorly. In illecehrosus the widest 
point lies about half-way between the insertion of the fins and the mantle 
opening; the body is more full, robust. 

Male oxygonius have a sharp triangular dorsal lobe at the mantle open- 
ing. While small lobes may be present in some specimens of other species, 
they are not as distinct. 



316 Proceedings of the Biological Society of Washington 

Thickness of the mantle wall is a difficult character to quantify, par- 
ticularly because of changes that may occur after spawning, or at time of 
fixation, but the mantle wall of oxygonius appears to be thin in relation 
to the mantle walls of other species. 

Fin angle. The angle of the fins in oxygonius is acute, each fin form- 
ing an angle of primarily 25°-35°, occasionally up to 40°, with the longi- 
tudinal axis of the mantle. The fin angle in illecehrosus is between 40°- 
50°, while in coindetii it generally exceeds 50°. Three specimens of 
argentinus have fin angles of 42°, 47° (females), and 50° (male). The 
ratio of fin length to fin width is 1/1.1 in oxygonius, 1/1.3 in illecehrosus, 
1/1.5 in coindetii, and 1/1.4 in argentinus. 

Hectocotylus. Hectocotyli of the species of Illex show the same basic 
structure; the tip of either right or left arm IV is modified in a series of 
lamellae and knobs (pi. 9). The proportional length of the modified 
section of the arm varies among the four species : argentinus, greater than 
50 percent (67 percent) of arm length; oxygonius, 30-35 percent; coin- 
detii, 30-35 percent; illecehrosus, 15-25 percent. The hectocotylus of 
coindetii, which is otherwise generally similar to that of oxygonius, has 
lobate trabeculae that are fringed and papillose (pi. 9C, D); this feature 
is lacking also in illecehrosus and argentinus. In coindetii from African 
waters the modified trabeculae are larger, more papillose than those on 
specimens from the Gulf of Mexico and the Mediterranean. The size of the 
lamellae along the dorsal row of the modified tip varies; oxygonius has 
the largest lamellae, coindetii and argentinus slightly smaller, and illece- 
hrosus smallest. A few knobs or papillae occur along the dorsal row prox- 
imal to the fully developed lamellae; oxygonius has about 3 knobs and 2 
slightly flattened papillae, illecehrosus and coindetii have 1-2 knobs. 
Illex argentinus has the most distinctive hectocotylus ( pi. 9B ) ; dorsal and 
ventral rows with 8-9 normal suckers pro"^imally; dorsal row continues 
with 16 enlarged, rounded, suckerless knobs, 18 large distinct lamellae 
and 6 or more small indistinct lamellae to the tip; ventral row continues 
with 7 low, suckerless knobs, 8 nipplelike papillae, and 20 or more small, 
indistinct, narrow lamellae to the tip. 

Spermatophores. The primary difference in the spermatophores of the 
four species occurs at the oral end of the cement body and the aboral end 
of the ejaculatory apparatus (pi. 10). In oxygonius (pi. lOA) the cone is 
funnel shaped and about equal-sided in profile with rounded corners; the 
oral tube is relatively broad; the aboral neck is intermediate in width and 
length in comparison with illecehrosus and coindetii. In illecehrosus the 
cone in profile is a low right-isosceles triangle with rounded corners; the 
oral tube is narrow; the neck is long and narrow (pi. lOB). In coindetii 
the cone is lens shaped or a rounded triangle or diamond shape in profile; 
the oral tube is broad; the neck is very short, broad, indistinct ( pi. IOC ) . 
In argentinus the cone is flat, lens shaped; the oral tube is broad; the 
neck is broad and distinct ( pi. lOD ) . 

Size and maturity. In species of Illex where sufficient material is 



The genus Illex 317 

available (e.g., illecehrosus, coindetii and probably argentinus) , the fe- 
males attain a larger maximum size than the males, and, in general, the 
males mature at a smaller size than the females. Males of illecehrosus in 
Newfoundland waters attain 2.70 mm ML but become sexually mature 
between 220-250 mm ML; females reach 310 mm ML but no fully ma- 
ture specimens have been observed there. In warmer waters illecehrosus 
does not grow as large as in cold waters and it reaches maturity at a 
smaller size. Illex coindetii attains a smaller maximum size than illece- 
hrosus and reaches maturity at a correspondingly smaller size. Throughout 
its geographic range, however, coindetii shows variation both in maximum 
size and in size at maturity. 

Studies now in progress on Illex, as well as the present analysis of 
characters, lead to the conclusion that the subspecific designations can 
no longer be justified. Based on the accumulated evidence, we consider 
that the differences among illecehrosus, coindetii, argentinus and oxy gen- 
ius are significant at the species level and we elevate these taxa to the 
status of full species. 

ARTIFICIAL KEY TO THE SPECIES OF ILLEX 

( This key is based on mature specimens. ) 

1. Hectocotylus greater than 50 percent of total arm length 



I. argentinus ( Castellanos, 1960) 

Hectocotylus less than 40 percent of total arm length 2 

2. Hectocotylus about 15-25 percent of total arm length; suckers on 

arms II and III of males not enlarged; head narrow in adult 

males I. illecehrosus (Lesueur, 1821) 

Hectocotylus about 30'-35 percent of total arm length; suckers on 
arms II and III of males enlarged; head broad in adult males 
3 

3. Fin angle 25°-40°; fin length/fin width index = 1/1.1; cone at oral 

end of cement body of spermatophore funnel shaped, length 

of neck about equal to width 7. oxygonius, n. sp. 

Fin angle greater than 45°; fin length/fin width index = 1/1.5; 
cone at oral end of cement body of spermatophore lens shaped, 
length of neck much less than width /. coindetii ( Verany, 1837 ) 

Distribution: Illex oxygonius has been captured at localities that range 
from about 24° to 39° N and from 73° to 84° W (fig. 1). This area in- 
cludes southern localities in the Gulf of Mexico, the Straits of Florida, 
and in the region of the Chesapeake Bi^ht (New Jersey to Virginia). The 
species is sympatric in at least parts of its range with several other species 
of Illex. It overlaps the southern portion of the range of illecehrosus, 
while the southern part of the range of oxygonius overlaps with the north- 
ern portion of the range of coindetii. 

Illex argentinus, so far as known, occurs along the Argentine coast 
from about 35° to 47° S and from 52° to 62° W (Castellanos, 1964; 



318 Proceedings of the Biological Society of Washington 

Castellanos and Menni, 1968). Undoubtedly, the species has a broader 
distribution than is presently recorded, but the proper delineation of its 
limits await further investigation based on additional collections. 

Illex illecehrosus is pan-Atlantic in far northern waters from New- 
foundland and Labrador (Grimpe, 1925), Greenland (Muus, 1962), Ice- 
land (Bruun, 1945), and northern European waters south to the Bristol 
Channel (Adam, 1952). In the western Atlantic illecehrosus extends 
southward from Newfoundland and Labrador all along the east coast of 
North America. Past records of illecehrosus from the Gulf of Mexico and 
Caribbean Sea very possibly are a combination of two species, coindetii 
and oxygonius, so can no longer be used as an indication of true species 
distribution. 

Specimens of illecehrosus currently available to us come from New- 
foundland and Labrador, Gulf of Maine, and along the east coast of the 
U.S. Among the southernmost records are the following specimens and 
localities : 

Oregon 6800, 7 males, 153-175 mm ML, 8 females, 155-202 mm ML, 

29°48'N 8O°09.5'W, 333 m 
Oregon 6801, 8 males, 148-170 mm ML, 2 females, 145-205 mm ML, 

29°39.5'N 80°08.5'W, 383 m 

Illex coindetii in the eastern Atlantic extends from reportedly the North 
Sea, certainly along the coasts of Portugal and west Africa to 14° S 
(Adam, 1952); it is very common in the Mediterranean (Mangold, 1963) 
and Adriatic Sea ( Gamulin-Brida and Ilijanic, in press). Now coindetii 
is reported for the first time from the western Atlantic waters of the Carib- 
bean Sea and the Gulf of Mexico. Among the specimens that we have 
verified as coindetii from this region are the following: 

Oregon II 289, 1 male, 135 mm ML, 11°24'N 73°47'W, 273 m 
Oregon 445, 4 males, 147-157 mm ML, 2 females, 211-217 mm ML, 

19°48'N91°20'W, 25 m 
Oregon 550, 2 males, 72-81 mm ML, 1 female, 162 mm ML, 26°55'N 

96°25.5'W, 228 m 
Oregon 3634, 1 male, 160 mm ML, 1 female, 215 mm ML, 16°44'N 

87°55'W, 346 m 
Oregon 4410, 1 male, 155 mm ML, 11°52'N 69°27'W, 420 m 
Oregon 4411, 1 male, 163 mm ML, 11°55'N 69°27'W, 473 m 
Oregon 4610, 1 male, 160 mm ML, 27°37'N 93°22'W, 364-382 m 
Triton Station, 1 male, 110 mm ML, 1 female, 145 mm ML, South of 

Palm Beach, Florida, in Florida Current. 
Tortugas, Florida, 1 female, 173 mm ML, Gulf of Mexico. 

Illex illecehrosus is predominantly a cold water species, but its exten- 
sion into southern waters indicates that it may be eurythermal. Although 
few experimental data are available. Frost and Thompson ( 1932, 1933, 
1934) have shown through an analysis of fisheries records that illece- 
hrosus in Newfoundland waters tolerates a temperature range of 0°-15° C, 



The genus Illex 319 

but optimal temperatures for maximum catches were 7°-15°; Squires' 
(1957) data confirm this temperature range. In southern waters illece- 
brosus must survive warmer temperatures if it occurs at shallow depths; 
however, our southernmost records, O 6800 and O 6801, are from 333-383 
m where the temperature is 9°-10°C. The vertical ranges of the western 
Atlantic species of Illex have yet to be determined, consequently little can 
be stated about temperature and depth relationships. 

Illex coindetii in the western Mediterranean ranges from about 40-500 
m and meets temperatures of 12.5°-20°C (Mangold, 1963). In West 
African waters coindetii was collected between 50-500 m where tempera- 
tures ranged from 7.8°-22.7°C (Adam, 1952). Generally similar tem- 
peratures occur in the western Atlantic range of coindetii. 

Illex argentinus is captured at depths of 7-800 m where temperatures 
range from 5°-12°C ( Castellanos, 1964; Castellanos and Menni, 1968). 

Illex oxygonius is recorded from depths of 50-555 m and from tempera- 
tures of 7°-13°C. 

The complexities of the Illex species systematics and distributions that 
have been discovered during the present study, particularly in the Carib- 
bean, Gulf of Mexico and Atlantic waters from Florida to the Chesapeake 
Bight, make it difficult at present to establish with certainly the south- 
ern limits of the ranges of illecehrosus, coindetii and oxygonius and the 
northern limits of coindetii and oxygonius. Examination of existing col- 
lections and systematic sampling for additional material, particularly from 
western Atlantic warm waters, are required to resolve the systematic and 
zoogeographical problems that presently preclude a complete understand- 
ing of the polytypic genus Illex. 

Literature Cited 
Adam, W. 1939. Notes sur les cephalopodes. 12. Illex illecehrosus 

( Lesueur ) trouve dans I'estomac d'Orthagoriscus mala Linne. 

Bull Mus. r. Hist. nat. Belg., 15: 1-3. 
. 1952. Cephalopodes. Result, sci. Exped. oceanogr. Belgium 

(1948-1949), 3(3): 1-142. 
Bruun, a. F. 1945. Cephalopoda. Zool. Iceland, 4(64): 1-15. 
Castellanos, Zulma J. A. de. 1960. Una nueva especia de calamar 

argentine Ommastrephes argentinus sp. nov. (MoUusca, 

Cephalopoda). Neotropica, 6 ( 20 ) : 55-58. 
. 1964. Contribucion al conocimiento biologico del calamar 

argentino Illex illecehrosus argentinus. Bol. Inst. Biol. Mar. 

Mar del Plata, 8: 1-37. 
. and R. C. Menni. 1968. Los cefalopodos de la expedicion 

"Walther Herwig." Notas Com. Invest. Cient. Bs. As., 6(2): 

1-31. 
Clarke, M. R. 1966. A review of the systematics and ecology of oceanic 

squids. Adv. mar. Biol, 4: 91-300. 
Degner, E. 1925. Cephalopoda. Rep. Danish oceanogr. Exped. Medi- 

terr., 2: 1-94. 



320 Proceedings of the Biological Society of Washington 

Frost, Nancy and H. Thompson. 1932. The squids. Ann. Rept. New- 
foundland Fish. Comm., 1(4): 25-33. 
. 1933. The squids. Ann. Rept. Newfoundland Fish. Comm., 

2(1): 80-86. 
. 1934. Squid. Ann. Rept. Newfoundland Fish. Comm., 2(2): 

59-64. 
Gamulin-Brida, Helena and Vesna Ilijanic. Quelques renseigne- 

ments sur la distribution des Cephalopodes lUicinae au large 

de I'Adriatique. Rapp. Proc.-Verb. Reun. C.I.E.S.M., 21 (in 

press ) . 
Grimpe, G. 19'2i1. Teuthologische Mitteilungen. VII. Systematische Ueber- 

sicht der Nordseecephalopoden. Zool. Anz., 52: 297-305. 
. 1924. Teuthologische Mitteilungen. IX-XI. Zool. Anz., 53: 

320-330. 
. 1925. Zur Kenntnis der Cephalopodenfauna der Nordsee. 

Wiss. Meeresunters Kiel, 16(3): 1-124. 
. 1933. Die Cephalopoden des arctischen Gebietes. Fauna 

Arctica, 6(5): 489-514. 
Lesueur, C. a. 1821. Descriptions of several new species of cuttlefish. 

Jour. Acad. Nat. Sci. Phila. 2: 86-101. 
Mangold-Wirz, Katharina. 1963. Biologie des cephalopodes benthi- 

ques et nectoniques de la Mer Catalane. Vie Milieu, Supple- 
ment No. 13, 385 pp. 
Mangold, Katharina, C. C. Lu and F. A. Aldrich. A reconsideration of 

forms of squid of the genus Illex ( lUicinae, Ommastrephidae ) . 

II. Sexual Dimorphism. Can. J. Zool. (In press). 
Muss, B. J. 1962. Cephalopoda. Meddelelser om Gr0nland, 81(5): 1- 

23. 
Naef, a. 1916. Ueber neue Sepioliden aus dem Golf von Neapel. Fubbl. 

Staz. Zool. Napoli, 1: 1-10. 
. 1921. Das System der dibranchiaten Cephalopoden und die 

Mediterranean Arten derselben. Mitt. Zool. Stat. Neapel, 22: 

527-542. 
. 1923. Die Cephalopoden. Systematic. Fauna u Flora Neapel, 

Monograph 35, 1 (pt. 1, no. 2): i-xiv, 149-863. 
Pfeffer, G. 1912. Die Cephalopoden der Plankton-Expedition. Ergebn. 

Planktonexped. Humboldt-Stiftung, 2:i-xxi, 1-815. 
SQuraES, H. J. 1957. Squid, Illex illecebrosus (Lesueur) in the New- 
foundland fishing area. Jour. Fish. Res. Bd. Canada, 14(5): 

693-728. 
Steenstrup, J. 1880. The interrelationships of the Ommastrephes-like 

cephalopods. An orientation. Overs, danske Vidensk. Selsk 

Forh. 1880-1881: 73-110. English translation: A. Volse, J. 

Knudsen and W. J. Rees. 1962. The cephalopod papers of 

Japetus Steenstrup. Danish Science Press. Copenhagen. 330 

pp. 



The genus Illex 321 

Verany, J. B. 1837. Memoire sur six nouvelles especes de Cephalopodes 
trouvees dans la Mediterranee a Nice. Mem. Ace. Sci. Torino, 
Ser. 2, 1: 91-98. 

Voss, G. L. 1956. A review of the cephalopods of the GiJf of Mexico. 
Bull. Mar. Sci. Gulf & Caribb., 6(2): 85-178. 

. 1962. List of the types and species of cephalopods in the 

collections of the Academy of Natural Sciences of Philadel- 
phia. Notulae Naturae, (365): 1-7. 

. 1963. Cephalopods of the Philippine Islands. United States 

Nat. Mus. Bull. 234: 1-180. 



322 Proceedings of the Biological Society of Washington 



^ QH ^ ^ 

1 

I B4X 

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J28 3 October 1969 

PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



A NEW CLINID FISH SPECIES, PARACLINUS 
FEHLMANNI, FROM ECUADOR 



By Victor G. Springer and Robert E. Tr] 
Smithsonkin Institution, Washington, D. 




0CT201969 



While routinely identifying fishes collected by Dr 
Fehlmann on the coast of Ecuador, we encountered a new spe- 
cies of Paraclinus Mocquard. This new species is the second in 
the genus known from Ecuador and it and its congener, P. mex- 
icanus, are the southernmost representatives of Paraclinus in 
the eastern Pacific. The new species represents the twelfth 
eastern Pacific member of Paraclinus, which was recently re- 
vised for that area by Rosenblatt and Parr ( 1969) . 

The methods used are those of Hubbs ( 1952 ) , except that all 
anal fin elements are counted (the last two anal rays were 
counted as two) and the caudal fin counts are of the number 
of segmented rays. 

Paraclinus fehlmanni new species 
Fig. 1 

Holotype: U.S. National Museum 202697, male, 40.3 mm standard 
length (SL), Bahia de Santa Elena, Ecuador (02° 13' 11" S, 80° 55' 22" 
W), area exposed by rocks at low tide and small tide pools, 0-6 feet, 
poisoned, 4 May 1966, H. A. Fehlmann, HA-102. 

Paratypes: USNM 202714, 13 (26.2-43.6), taken with the holotype, 
and 202713, 3 (17.1-22.7), tip of Punta Ancon, Anconcito village, Golfo 
de Guayaquil, Ecuador (02° 20' 33" S, 80° 53' 40" W), large tide pool 
formed by sandstone, both cemented and loose rocks, 0-5 feet, 5 May 
1966, H. A. Fehlmann, HA-103. 

Description: (Asterisks signify characters of the holotype.) Dorsal fin 
consisting of spinous elements only: XXIX (3 specimens), XXX(13)*, 
XXXI(l); anal fin spines II*, rays 18 (8), 19(9)*; pectoral rays 13 
(16)*, 14(1); pelvic rays I, 3*, innermost ray separated from middle ray 
by a well-developed membrane*; segmented caudal rays 13*; branchios- 

23— Proc. Biol. Soc. Wash., Vol. 82, 1969 (323) 



324 Proceedings of the Biological Society of Washington 



,r 




^^^^' 






New Ecuador fish 325 

tegal rays 6*; precaudal vertebrae 11*, caudal vertebrae 24 (3) or 25 
(14)*, total, 35 or 36*; lateral line scales in arched portion 15 (1), 16 
(15)*, 17 (1); in straight portion 19 (7)*, 20 (10), total, 35 (8)* or 
36 (9). 

All scales without flaplike projections on posterior margins. Rosenblatt 
and Parr ( 1969, fig. 1 ) use the nature of squamation in the region behind 
the nape and above the anterior end of the lateral line as a specific 
character in Paraclinus. Specimens of P. fehlamnni exhibit all grada- 
tions from a large, naked area (their fig. la) to a restricted naked area 
(their fig. lb) in this region. Pectoral base naked. Opercular spine 
simple, flat (not needlelike), rounded to pointed triangle, extending pos- 
teriorly to level of second to third dorsal spine. Nuchal and orbital cirri 
palmate, irregularly frayed or with a few equal branches on distal mar- 
gins ( both cirri simple flaps in one specimen ) . Nasal cirrus, simple, short 
filament. Dorsal fin membrane incised not at all to two-fifths (one- 
eighth or less in 11 of 17 specimens) length of fourth dorsal spine, be- 
tween third and fourth dorsal spines. 

Proportional measurements, as percent SL, are given here for the 14 
specimens ranging from 26.2-43.6 mm SL. Except for orbital diameter, 
lengths of upper jaw, preanal region, and first and second dorsal spines 
(all discussed below) there are no noticeable sexual or size associated 
differences in proportions in these specimens. The proportions are given 
as ranges followed in parentheses by averages. 

Head length, 24.6-28.0 (26.1); snout length, 5.4-7.2 (6.0); inter- 
orbital width, 2.1-3.7 (3.1); predorsal length, 20.6-22.6 (21.9); origin 
of first dorsal spine to insertion of last spine, 74.0-79.4 (76.2); origin of 
first anal spine to insertion of last ray, 41.0-47.7 (44.6); caudal peduncle 
depth, 6.9-8.8 (7.6); horizontal preopercle width, 6.0-8.7 (6.8); inser- 
tion of first dorsal spine to origin of third, 3.6-5.5 (4.9); insertion of 
third dorsal spine to origin of fourth, 1.5-5.1 (4.0); length of third dorsal 
spine, 6.1-9.8 (8.2); length of fourth dorsal spine, 5.3-7.4 (6.5); length 
of 22nd dorsal spine, 8.4-10.5 (9.2); length of penultimate dorsal spine, 
5.5-8.1 (7.0); length of ultimate dorsal spine, 4.3-7.2 (5.5); length of 
first anal spine, 4.2-6.0 (5.4); length of second anal spine, 5.4-7.2 (6.4); 
length of longest caudal ray, 19.5-22.6 (20.9); length of longest pectoral 
ray, 19.2-22.8 (20.7); length of longest pelvic ray, 14.2-18.6 (16.3); 
nuchal cirrus length 1.2-2.2 (1.6); nasal cirrus length, 0.8-1.8 (1.2); 
orbital cirrus length, 1.3-2.7 (1.9). 

Orbital diameters gave evidence of relative decrease with increase in 
SL. In three specimens 17.1-22.7 mm SL, they were 7.6-8.9; in two 
specimens 26.2-29.4, they were 7.3-8.0; in nine specimens 31.9-37.4, 
they were 6.4-7.5; in three specimens 40.3-43.6, they were 5.4-7.2. 

Upper jaw lengths in the six males, 12.5-17.9 ( 15.0); jaw extending to 
or beyond level of posterior margin of orbit (four males), to slightly 
beyond mid-orbital level (two males). Upper jaw lengths in the eight 
females, 11.5-13.1 (12.1); jaw extending to level of mid-orbit in three 






326 Proceedings of the Biological Society of Washington 

females, slightly beyond mid-orbit in three females, to posterior margin 
of orbit in two females. 

Preanal length in the six males, 49.3-51.8 (50.8), apparently not 
changing relative to size; in the eight females, 50.8-56.9 (53.7), in- 
creasing in relative size with increase in SL. 

First dorsal spine length in five males, 9.4-12.2 (10.7), possibly in- 
creasing in relative length with increase in SL; in seven females, 11.2- 
14.7 (13.0), increasing in relative length with increase in SL; no overlap 
in relative spine lengths between males and females of similar SL. 

Second dorsal spine lengths in four males, 10.0-11.4 (10.4); no indica- 
tion of increase in relative length with increase in SL; in eight females, 
10.7-14,3 (12.6); increasing in relative length with increase in SL; no 
overlap in spine lengths between males and females of similar SL. 

Color in alcohol: Head and body varying from completely pale to ir- 
regularly dusky, with posterior scale margins outlined in dusky. No evi- 
dence of banding on body in any specimen. Dusky specimens with a pale 
to dusky, darkly outlined bar or line extending posteroventrally from or- 
bit. Dorsal, anal, and caudal fins clear to dark dusky; dark spot over 
21st to 23rd dorsal spine in 13 of 17 specimens ( absent in 4). Caudal oc- 
casionally with some evidence of alternating dark and pale spots on rays. 
Pectorals and pelvic rays with alternating dark and clear spots in all 
specimens. 

Etymology: Named for H. Adair Fehlmann, Smithsonian Oceano- 
graphic Sorting Center, in recognition of his outstanding contributions to 
ichthyology as evidenced by a long and productive history of ichthyologi- 
cal collecting. 

Discussion: Some specimens of Paraclinus fehlmanni will key to state- 
ment 2a, P. beebei Hubbs, in Rosenblatt and Parr's (1969) key to 
eastern Pacific Paraclinus. Other specimens will not key beyond state- 
ment lb, and are thus unidentifiable with any other species of Paraclinus. 
In lacking scales on the pectoral base, P. fehlmanni differs from all other 
eastern Pacific species of Paraclinus except P. beebei, to which it appears 
to be most closely related. It differs from P. beebei, which is known from 
Baja California to Panama, in having: spotted pectoral fins (unmarked in 
P. beebei, R. H. Rosenblatt, in litt.), 29 to 31 dorsal spines (usually 30; 
27 to 29 in P. beebei); 18 or 19 segmented anal rays (16 to 18 in P. 
beebei); 35 or 36 lateral line scales (32 to 35 in P. beebei); and in com- 
monly attaining a size over 36 mm SL (rarely to 36 mm in P. beebei; 
most specimens under 30 mm). 

Aside from lacking scales on the pectoral base, P. fehlmanni can be 
separated from its other eastern Pacific congeners as follows ( only some 
differences noted ) : from P. sini Hubbs in lacking membranous flaps on 
the posterior scale margins; from P. ditrichus Rosenblatt and Parr in hav- 
ing I, 3 pelvic rays (as opposed to I, 2); from P. mexicanus (Gilbert), 
also known from Ecuador, P. magdalanae Rosenblatt and Parr, P. ste- 
phensi Rosenblatt and Parr, and P. monophthalmus ( Giinther ) in lacking a 



New Ecuador fish 327 

segmented dorsal ray; from P. walkeri Hubbs in having 6 branchiostegal 
rays ( instead of 7 ) ; from P. integripinnis ( Smith ) in having the oper- 
cular spine always simple and in usually having fewer dorsal spines and 
anal rays; from P. tanygnathus Rosenblatt and Parr in having one to three 
more dorsal spines, two or three more lateral line scales, and in not hav- 
ing the membrane between the third and fourth dorsal spines deeply 
notched; and from P. altivelis ( Lockington ) in lacking a segmented dorsal 
ray and in having the first dorsal spine much shorter than the predorsal 
length. 

Pa/racUnus fehlmanni differs from all it Atlantic congeners in having 
the combination of three segmented pelvic rays with no segmented dor- 
sal ray. 

Rosenblatt and Parr reaffirmed Hubbs ( 1952 ) and Springer's ( 1955 ) 
findings that overall phyletic lines are not possible to distinguish in 
Paraclinus. The additional new species contributes nothing more to such 
discussion. 

Literature Cited 

Hubbs, C. 1962. A contribution to the classification of the blennioid 
fishes of the family Clinidae with a partial revision of the 
eastern Pacific forms. Stanford Ichthyol. Bull., vol. 4, no. 2, 
pp. 41-165. 

Rosenblatt, R. H. and T. D. Parr. 1969. The Pacific species of the 
clinid fish genus Paraclinus. Copeia, 1969, no. 1, pp. 1-20. 

Springer, V. G. 1955. Western Atlantic fishes of the genus Paraclinus. 
Texas Journ. Sci. (1954), vol. 6, no. 4, pp. 422-441. 



328 Proceedings of the Biological Society of Washington 



I 



QH 7^ 

1 

B4X 

HH 

348 3 October 1969 

PROCEEDINGS /^"^^^fev 

OF THE / ^^ 

BIOLOGICAL SOCIETY OF WASHINGTQNT^' 2 

TWO NEW SPECIES OF THE CRAYFISH GENUS 

PROCAMBARUS ( DECAPODA, ASTACIDAE ) WITH KEYS 

TO THE MEMBERS OF THE SPICULIFER GROUP 

By Horton H. Hobbs, Jr. 
Smithsonian Institution, Washington, D.C. 

The two species described here have Hmited ranges in Louisi- 
ana and Georgia. That from Louisiana has been found in a few 
localities in the Ouachita drainage system in Claiborne, Ouach- 
ita, and Union parishes, and that from Georgia in the Flint 
River System in Crawford, Lee, Schley, and Sumter counties. 

Keys to the species of the Spiculifer Group, one based on 
first form males and the other on females, are provided and 
include summaries of the ranges of each. 

Procambarus elegans new species 
(Figures 1-11, 24, 25) 

Diagnosis: Body pigmented, eyes well-developed. Rostrum with gently 
convergent margins bearing pair of marginal spines; acumen long and 
slender. Areola 5.19 to 5.9 times longer than wide and constituting 26.6 
to 28.6 percent o£ entire length of carapace. Carapace with two cervical 
spines on each side. Suborbital angle small and rounded. Postorbital ridges 
terminating in spines. Antennal scale approximately 2.5 times longer 
than wide, broadest proximal to midlength. Mesial margin of palm of 
chela with seven or eight tubercles, and both fingers provided with 
moderately well defined longitudinal ridges. Ischiopodites of third and 
fourth pairs of pereiopods with hooks, that of fourth weakly bituber- 
culate; coxae of fourth and fifth pereiopods with prominences. First 
pleopods asymmetrical, without shoulder on cephalic surface, reaching 
cephalad to coxae of third pair of pereiopods, and provided with subter- 
minal setae; distal extremity bearing (1) slender acute mesial process 
directed mesiodistally and slightly caudally, (2) short spinelike cephalic 
process arising from cephalic surface of appendage and extending cephalo- 
distally, (3) caudal element consisting of truncate caudal knob bearing 
minute, acute, toothlike caudal process directed cephalodistally, and (4) 
short, corneous central projection, situated immediately caudal to cephalic 

24— Proc. Biol. Soc. Wash., Vol. 82, 1969 (329) 



330 Proceedings of the Biological Society of Washington 




Figs. 1-4. Procambarus elegans new species. 1, Dorsal view of holo- 
type. 2, Mesial view of first left pleopod of holotype. 3. Annulus ventralis 
of allotype. 4, Lateral view of first left pleopod of holotype. 



Two new crayfish 331 

process, extending subparallel to it, and reaching distally to approximately 
same level; neither cephalic process nor central projection visible in 
caudal aspect of appendage. 

Holotypic Male, Form I: Body subovate, appearing slightly compressed. 
Abdomen narrower than thorax (18.3 and 23 mm). Width of carapace 
slightly greater than height at caudodorsal margin of cervical groove 
(23.0 and 22.2 mm). Areola 5.9 times longer than wide with four or 
five punctations across narrowest part. Cephalic section of carapace 2.3 
times as long as areola (length 28.2 percent of entire length of carapace). 
Rostrum excavate dorsally with unthickened margins parallel at base and 
convergent distally; upper surface of rostrum with usual submarginal setif- 
erous punctations and others scattered between; well-developed marginal 
spines present at base of acumen; acumen long and spiniform, constitut- 
ing approximately 42 percent of entire length of rostrum. Subrostral 
ridges weak and visible in dorsal aspect only near base of rostrum. Post- 
orbital ridges moderately prominent, grooved dorsally and terminating 
cephalically in prominent spines. Suborbital angle weak, almost obsolete. 
Branchiostegal spine acute and prominent. Carapace punctate dorsally, 
tuberculate cephalolaterally and granulate ventrolaterally; row of tuber- 
cles along ventral margin of cephahc portion of cervical groove. Two 
cervical spines present on each side of carapace, more ventral ones larger. 
Abdomen subequal in length to carapace. Cephalic section of telson 
with three spines in each caudolateral corner. Epistome (Fig. 8) sub- 
equal in length and breadth; margin shghtly elevated and converging to 
subacute apex. Antennules of usual form with prominent spine on ventral 
surface of basal segment near midlength. Antennae extending caudad 
to fourth abdominal tergum. Antennal scale (Fig. 7) about 2.7 times 
longer than broad, broadest proximal to midlength with widest lamellar 
area about 1.6 times width of thickened lateral portion, latter terminating 
in strong acute spine. 

Right chela (Fig. 1) depressed with palm somewhat inflated; lateral 
margin with row of tubercles extending to base of costate distal third of 
fixed finger; upper and lower surfaces of palm tuberculate with mesial 
row of eight tubercles subtended dorsolaterally by irregular row of seven; 
lower surface of palm with prominent tubercle lateral to articular condyle 
at base of dactyl. Fixed finger with moderately well-defined submedian 
ridges flanked dorsally by squamous tubercles basally and setiferous 
punctations distally; opposable margin with row of 17 tubercles (sixth 
from base largest) along basal three-fourths with yet larger tubercle be- 
low former row at base of distal third, and crowded minute denticles 
present along almost entire length below upper row of tubercles; lower 
surface with poorly defined submedian ridge flanked along most of its 
length by setiferous punctations and proximally by few scattered tuber- 
cles; ventromesial surface with row of six tubercles along proximal half 
of finger. Dactyl broken and partially regenerated but in paratypic first 
form male bearing weak submedian ridge dorsally and ventrally, both 



5»jnH^nrnaK nn-rS 1989 



332 Proceedings of the Biological Society of Washington 




Figs. 5-11. Procambarus elegans new species (pubescence removed 
from all structures illustrated). 5, Caudal view of first left pleopod of 
holotype. 6, Mesial view of first left pleopod of holotype. 7, Antennal 
scale of paratypic male, form I. 8, Epistome of paratype. 9, Lateral view 
of first left pleopod of holotype. 10, Basal portions of third and fourth 
pereiopods of paratypic male, form I. 11, Lateral view of carapace of 
holotype. 



flanked by tubercles proximally and setiferous punctations distally; 
lateral surface subserrate with row of tubercles extending almost to apex; 
opposable margin convex laterally with greatest height of convexity just 
proximal to midlength, with upper row of 13 small rounded tubercles 
along proximal two-thirds and row of four distinctly larger ones along 
antipenultimate fourth; most proximal one in latter row so large as to give 
impression of excision on margin immediately proximal to it; crowded 
minute denticles present between and distal to tubercles and between 
rows. 

Carpus of right cheliped longer than broad (14.1 and 8.9 mm), with 
mesial portion tuberculate and area lateral to dorsal diagonal excavation 



lolotype 


Allotype 


Morphotype 


22.2 


13.2 


14.6 


23.0 


13.9 


15.0 


53.2 


31.6 


37.5 


2.6 


1.6 


1.8 


15.2 


8.3 


10.2 


8.8 


5.0 


5.7 


17.8 


11.3 


12.9 


16.6 


4.9 


6.6 


14.7 


4.5 


6.0 


46.4 


14.6 


20.5 


broken 


8.3 


12.9 



Two new crayfish 333 

Table 1. Measurements (mm) of Procambarus elegans. 



Carapace: 
Height 
Width 
Length 

Areola: 
Width 
Length 

Rostrum : 
Width 
Length 

Chela: 

Length of inner margin of palm 
Width of palm 

Length of outer margin of chela 
Length of dactyl 



with setiferous punctations; distal dorsomesial surface with prominent 
spine; mesial surface with one spiniform tubercle near midlength and 
several smaller ones proximal to it; lower surface with two distal spines, 
lateral one on ventral articular condyle and other more mesially situated; 
six small tubercles situated proximomesial to mesial spine. 

Merus of right cheliped tuberculate dorsally, distomesially, and ven- 
trally, otherwise punctate; upper distal surface with two prominent 
spines; ventral surface with irregular lateral row of seven tubercles, some 
small, others spikelike, and more regular lateral row of 12, generally 
larger distally; small tubercles present between two rows, and row of 
three connecting two rows distally. Ischium with mesial row of three tu- 
bercles. 

Hooks on ischia of third and fourth pereiopods ( Fig. 10 ) ; that on third 
simple that on fourth weakly bituberculate. Coxa of fourth pereiopod 
with prominent rounded boss caudomesially; that of fifth with small 
slender projection lateral to penis papilla. 

Sternum between second, third, and fourth pereiopods moderately deep 
and bearing prominent fringe of setae on ventrolateral margins. 

First pleopods ( Figs. 2, 4-6, 9 ) as described in diagnosis. 

Allotypic Female: A juvenile female which differs in no important 
respects, other than secondary sexual characters, from holotype. Chela 
conspicuously shorter and smaller with minute denticles between fingers 
reduced to single rows. Annulus ventralis (Fig. 3) movable, with shallow 
median trough cephalically; sinus originating along median line almost at 
midlength of annulus, making C-shaped turn sinistrally, returning to 



334 Proceedings of the Biological Society of Washington 

median line and extending caudally almost to caudal margin. Sternum 
cephalic to annulus devoid of tubercles or caudally projecting lobes. 

Morphotypic Male, Form II: Differs from holotype in only few minor 
details: Cephalic section of telson with three spines in one caudolateral 
corner and two in other; mesial margin of palm of chela with only seven 
tubercles; fixed finger of chela with row of 10 tubercles, third from base 
largest; dactyl of chela with upper row of three tubercles and lower of 
five, proximal tubercle in lower row largest; merus of cheliped with 
lower lateral row of only five tubercles and inner one of 11; ischia of 
chelipeds with two tubercles on one and four on other; hooks on ischia of 
third and fourth pereiopods much reduced in size and with no indication 
of bitubercular condition on that of fourth; prominences on caudomesial 
surface of coxa of fourth pereiopod lower and more rounded. 

First pleopod (Figs. 24, 25) with all terminals inflated, but mesial and 
cephalic processes heavier, and central projection and caudal process 
much less distinctly delimited, than in holotype. 

Color Notes: Dorsum of carapace straw brown with pair of irregular 
dark brown longitudinal stripes extending from below postorbital ridges to 
caudal margin of carapace where broadened mesially to form two lobes 
(remnants of bar, see Hobbs 1958, p. 74, figs. 1 and 2) reaching dorsally 
to level of branchiocardiac grooves. Branchiostegites ventral to longi- 
tudinal stripes pinkish tan fading ventrally to cream cephalically and 
lavender along most of ventral border; pinkish tan area flecked with ir- 
regular dark brown blotches. Rostral and postorbital ridges light tan. 
Abdomen with pale straw median longitudinal stripe subtended in each 
somite by pair of dark brown oblique ( cephalomesial to caudolateral) 
stripes which together form irregular stripe flanking median light stripe; 
areas ventral to dark stripes pinkish tan with irregular dark brown blotches. 
Telson bordered in dark brown with pale tan central area marked 
by irregular dark brown lines and blotches; lateral ramus of uropod with 
irregular dark brown lateral and mesial borders, and dark brown distal 
segment surrounding cream tan central area; mesial ramus dark brown 
except proximomesially where cream tan with irregular dark brown spots. 
Sternum of cephalothoracic and abdominal regions pinkish cream. Dorsal 
surface of cheliped distal to ischium pale tan with black tubercles and 
dark brown blotches; distal spines on merus and mesial spines and tuber- 
cles on carpus, propodus and dactyl with white tips. Base of fixed finger 
reddish brown, that of dactyl tan; both fingers with broad blackish band 
preceding orange-colored tips; lower surface pinkish cream except for en- 
circling blackish bands on fingers. Remaining pereiopods, from ischium 
distally, with tan lateral surfaces bearing darker brown mottlings — other- 
wise pinkish cream. 

Type-locality : Spillway from Corney Lake, an impounded tributary of 
the Ouachita River, Claiborne Parish, Louisiana. There the "coffee- 
tinted" water flows with a sluggish to moderately swift current through 
multiple channels in an eroded clay substrate which supports scattered 



Two new crayfish 335 

groups of aquatic plants including Myriophyllum, Potomageton, and 
Ludwigia. The confluence of the channels forms a moderately flowing 
stream some 40 feet wide and more than five feet in depth. 

Disposition of types: The holotypic male, allotypic female, and mor- 
photypic male, together with a single paratypic first form male, are de- 
posited in the Smithsonian Institution, nos. 129892, 129893, 115838, 
129894, respectively. 

Size: Only two first form males are available, and the carapace of one 
of them is crushed; the holotype has a carapace length of 53.2 mm, larger 
than that of any other specimen available. 

Range, Specimens Examined, and Crayfish Associates: Procambarus 
elegans is known from only five localities in the Ouachita River, system in 
Louisiana: CLAIBORNE PARISH— Type-locality, 26 May 1952 (1 $ 
I) T. H. Nickersoji coll.; 24 April 1965 (1 ^ I, 1 juv. 2 ) W. J. Harman 
and Horton H. Hobbs, Jr. coll., with P. a. acutus (Girard, 1852: 91), P. 
clarkii (Girard, 1852: 91), P. vioscai Penn, 1946: 27, Camharellus puer 
Hobbs, 1945: 469, Fallicambarus hedgpethi (Hobbs, 1948: 224), and 
Faxonella clypeata (Hay, 1899: 122). OUACHITA PARISH— Choud- 
rant Bayou, 8 December 1964 (1 $ II) N. H. Douglas, coll. with P. 
vioscai and O. palmeri longimanus (Faxon, 1898: 655). 

UNION PARISH— Mill Creek, Sec. 36— T 19N— R IE, 18 June 1965 
( 1 juv. $ ) NHD, coll. with P. vioscai and Orconectes palmeri longi- 
manus. Meridian Creek, 1 mi. E of Conway, Sec. 7 — T 22N — R IE, 21 
May 1956 ( 1 juv. $) W. R. Taylor and L. W. Lowe, coll. Meridian 
Creek, Sec. 21— T 22N— R IE, 5 June 1965 (1 juv. $, 1 juv. 2 ) NHD, 
coll. with P. vioscai and P. clarkii. 

Relationships: Procambarus elegans resembles those members of the 
Spiculifer Group which have retained the cephalic process on the first 
pleopod (see Key to First Form Males) and probably has its closest af- 
finities with P. natchitochae Penn^ 1953, and P. ablusus Penn, 1963. It 
differs from P. natchitochae in possessing a much shorter central projec- 
tion and a truncate caudal knob on the first pleopod, and its areola is 
much narrower and longer. It differs from P. ablusus in that the truncate 
caudal knob of the first pleopod obscures the caudal process and central 
projection in caudal aspect, the cephalic process and central projection 
do not reach so far distally, and the latter is not recurved. 

Etymology: The name elegans (L.), choice or select, was chosen 
because of the coloration and apparent rarity of this crayfish. 

Remarks: The first specimen (the paratypic male) that I saw was 
donated to me by Lt, T. H. Nickerson who indicated that it had been 
caught on a fishing line at the spillway of Corney Lake in swift water four 
to five feet deep on 26 May 1952. Not until 1965 did I have an op- 
portunity to visit the locality. Dr. Walter J. Harman accompanied me 
there, and both of us spent some four hours attempting to secure ad- 
ditional specimens by the usual seining and dip-netting techniques. 
Although at least 100 crayfishes were caught (47 retained), only two of 



336 Proceedings of the Biological Society of Washington 




Two new crayfish 337 

them were P. elegans. Perhaps this species is more abundant in the 
deeper waters of the spillway but it was our impression that it was ex- 
ceedingly rare in the area immediately below the dam. The holotype was 
secluded under a large log that had become lodged against the bank of 
one of the larger channels. 

Procambarus gibbus new species 
(Figures 12-23) 

Procambarus spiculifer. — Hobbs, 1953b: 416 [map of distribution of P. 
spiculifer, in part]. 

Diagnosis: Body pigmented, eyes well-developed. Rostrum with gently 
convergent margins bearing pair of marginal spines; acumen long and 
slender. Areola 2.8 to 3.5 times longer than wide and constituting 23.8 to 
27.2 percent of entire length of carapace. Carapace with two cervical 
spines on each side. Suborbital angle small and rounded. Postorbital 
ridges terminating in spines. Antennal scale broadest proximal to mid- 
length. Mesial margin of palm of chela with five or six tubercles, and 
both fingers provided with moderately to poorly defined longitudinal 
ridges. Ischiopodites of third and fourth pairs of pereiopods with hooks, 
that of fourth weakly bituberculate; coxae of fourth and fifth pereiopods 
with prominences. First pleopods asymmetrical, without shoulder on 
cephalic surface, reaching cephalad to coxae of third pair of pereiopods 
and provided with subterminal setae; distal extremity lacking cephalic 
process but bearing ( 1 ) slender, acute, slightly curved mesial process di- 
rected caudolaterally and reaching scarcely beyond level of tip of central 
projection, (2) caudal element consisting of simple, small bladelike cor- 
neous process closely applied to caudal margin of central projection, mesial 
adventitious ridge, and caudal knob poorly delimited from much swollen 
caudolateral end of shaft but somewhat fingerlike at lateral base of caudal 
process, and (3) central projection corneous, very conspicuous, beaklike 
but extending almost directly distally even though cephalic margin con- 
vex. Annulus ventralis with median longitudinal groove between elevated 
lateral walls and with flat, subtriangular midcaudal depressed area 



Figs. 12-23. Procambarus gibbus new species (pubescence removed 
from structures illustrated). 12, Mesial view of first left pleopod of 
holotype. 13, Mesial view of first left pleopod of morphotype. 14, Dor- 
sal view of carapace of holotype. 15, Lateral view of first left pleopod 
of morphotype. 16, Lateral view of first left pleopod of holotype. 17, 
Antennal scale of paratypic male, form I. 18, Epistome of holotype. 19, 
Basal portions of third and fourth pereiopods of holotype. 20, Annulus 
ventrahs of allotype. 21, Caudal view of first left pleopod of holotype. 22, 
Lateral view of carapace of holotype. 23, Dorsal view of distal podomeres 
of cheliped of holotype. 



338 Proceedings of the Biological Society of Washington 

crossed by straight or obliquely directed sinus. Sternum of female im- 
mediately cephalic to annulus with pair of very small lobes extending 
caudally, otherwise unadorned. 

Holotypic Male, Form I: Body subovate, compressed. Abdomen nar- 
rower than thorax (14.8 and 16.2 mm). Width of carapace slightly less 
than height at caudodorsal margin of cervical groove ( 16.2 and 17.0 
mm). Areola 3.2 times longer than wide with seven or eight punctations 
across narrowest part. Cephalic section of carapace 3.0 times as long as 
areola (length 24.9 percent of entire length of carapace). Rostrum ex- 
cavate dorsally with unthickened margins parallel at base and convergent 
distally; upper surface with usual submarginal setiferous punctations and 
others scattered between; well-developed marginal spines present at base 
of acumen; acumen long and spiculiform, constituting approximately 40 
percent of entire length of rostrum. Subrostral ridges weak and not visi- 
ble in dorsal aspect. Postorbital ridges prominent, grooved dorsolaterally, 
and terminating cephalically in prominent spines. Suborbital angle weak, 
obtuse, and almost obsolete. Branchiostegal spine acute and moderately 
prominent. Carapace punctate dorsally, granulate laterally, and tuber- 
culate ventrolaterally with row of tubercles along ventral margin of ce- 
phalic portion of cervical groove. Two strong cervical spines present on 
each side of carapace, more ventral one slightly heavier. Abdomen sub- 
equal in length to carapace. Cephalic section of telson with two spines 
in each caudolateral corner. Epistome (Fig. 18) about 1.5 times broader 
than long with margins slightly elevated, undulating, and converging to 
acute apex. Antennules of usual form with prominent spine on ventral 
surface of basal segment slightly distal to midlength. Antennae extend- 
ing caudad slightly beyond caudal margin of telson. Antennal scale ( Fig. 
17) about three times longer than broad, broadest proximal to midlength 
with widest lamellar area about 1.8 times width of thickened lateral por- 
tion, latter terminating in strong acute spine. 

Right chela ( Fig. 23 ) somewhat depressed with palm slightly inflated; 
lateral margin with tubercles extending to midlength of fixed finger with 
those distal to base of finger low and not visible in position in which 
chela figured, margin not costate; upper and lower surfaces of palm 
tuberculate with mesial row of six tubercles, lower surface with prominent 
tubercle distolateral to articular condyle at base of dactyl. Fixed finger 
with moderately developed dorsal submedian longitudinal ridge flanked 
by squamous tubercles proximally and setiferous punctations distally; op- 
posable margin with row of 20 tubercles (fifth from base largest) along 
proximal five-sixths, distal eight tubercles quite small; three tubercles 
below former row at base of distal fourth of finger, middle of three larger 
than other tubercles on opposable margin; minute denticles present along 
almost entire length between and below upper row of tubercles, lower 
surface without submedian ridge but with tubercles and punctation ar- 
ranged much like those on upper surface. Dactyl with very low dorsal 
and ventral submedian longitudinal ridges flanked proximally by tuber- 



Two new crayfish 339 

cles and distally by punctations; lateral surface subserrate with row of 
tubercles extending almost to apex; opposable margin convex with great- 
est height of convexity slightly proximal to midlength and bearing single, 
slightly irregular, row of 19 tubercles ending at base of distal one-eighth 
of finger, eighth tubercle from base largest, minute denticles present be- 
tween proximal tubercles, below more distal ones, and continuing almost 
to base of corneous tip of finger. 

Carpus of right cheliped longer than broad (10.2 and 6.5 mm), with 
mesial and dorsomesial surfaces tuberculate and dorsolateral, lateral, and 
ventral surfaces mostly punctate; dorsolateral and mesioventral surfaces 
also with few tubercles; distal dorsomesial surface with prominent spine; 
mesial surface with one spiniform tubercle near midlength, several smaller 
ones proximal to it, and one slightly distal to base of large tubercle; lower 
distal margin with two spines, lateral one on ventrolateral articular con- 
dyle and other more mesially situated with group of seven smaller tuber- 
cles proximomesial to it. 

Merus of right cheliped tuberculate dorsally, distomesially and ven- 
trally, otherwise punctate; upper distal surface with two prominent spines; 
ventral surface with irregular lateral row of 11 tubercles, three of which 
spikelike, and more regular lateral row of 15 tubercles, generally larger 
distally; additional small tubercles flanking rows; rows joined distally by 
oblique row of three tubercles. Row of four tubercles on ischium cor- 
responding to mesial row on merus. 

Hooks on ischia of third and fourth pereiopods (Fig. 19); that on 
third simple, that on fourth weakly bituberculate. Coxa of fourth pereio- 
pod with prominent, vertically disposed, rounded boss caudomesially; 
that of fifth with prominent slender projection caudomesially. 

Sternum between second, third, and fourth pereiopods deep and bear- 
ing moderate fringe of setae on ventrolateral margins. 

First pleopods ( Figs. 12, 16, 21 ) as described in diagnosis. 

Allotypic Female: Except in secondary sexual characteristics, differing 
only in minor details from holotype: areola with nine or 10 punctations 
across narrowest part; epistome with more prominent and acute cephalo- 
median projection; opposable margin of fixed finger of chela with 11 tu- 
bercles in upper row (fourth from base largest) and two below with mi- 
nute denticles arranged in single row; opposable margin of dactyl of chela 
with row of 13 tubercles (fourth from base largest) along proximal two- 
thirds of finger; ventral surface of merus of chela with lateral row of eight 
tubercles and mesial one of 1 1 ( See measurements ) . 

Annulus ventralis ( Fig. 20 ) shallowly situated in sternum, broader than 
long, with conspicuous median longitudinal trough extending between 
elevated (ventrally) lateral walls. Sinus arising on median line just 
cephalic to midlength making C-shaped curve sinistrally around fossa, 
continuing dextrally across median line before traversing flat, triangular 
caudal shelf to midcaudal margin of annulus. Sternum cephalic to an- 
nulus not tuberculate but with pair of small caudally projecting lobes. 



340 Proceedings of the Biological Society of Washington 




Figs. 24-38. First sinistral pleopods of members of Spiculifer Group 
(pubescence not indicated): Figs. 24 and 32, mesial views; Figs. 25-31 
and 33-38, lateral views. 24 and 25, Morphotype of F. elegans. 26, P. 
ablusus. 27, P. natchitochae. 28, P. penni. 29, P. vioscai. 30, P. dupratzi. 
31, P. echinatus. 32 and 33, P. suttkusi. 34, P. versutus. 35, P. ouachitae. 
36, P. lagniappe. 37, P. raneyi. 38, P. spiculifer. 



Two new crayfish 341 

Table 2. Measurements (mm) of Procambarus gibbus. 









Holotype 


Allotype 


Morphotype 


Carapace : 












Height 






17.0 


18.9 


16.7 


Width 






16.2 


17.9 


15.7 


Length 






37.0 


4L4 


38.0 


Areola: 












Width 






2.9 


3.4 


3.0 


Length 






9.2 


10.0 


9.2 


Rostrum : 












Width 






5.9 


6.5 


6.2 


Length 






12.9 


15.2 


13.9 


Chela: 












Length of 


inner margin 


of palm 


13.0 


8.4 


7.9 


Width of 


palm 




10.1 


7.4 


5.6 


Length of 


outer margin 


of chela 


34.7 


23.7 


21.0 


Length of dactyl 




19.0 


14.2 


12.1 



Morphotypic Male, Form II: Like allotype, other than in secondary 
sexual characteristics, differs only in minor details from holotype: fixed 
finger of chela with upper row of nine tubercles (fourth from base lar- 
gest) and single large one below row; dactyl of chela with row of 13 tu- 
bercles (fifth from base largest); ventral surface of merus of chela with 
lateral row of 11 tubercles and mesial one of 13; ischium of right chela 
with one spine and one tubercle; hooks on ischia of third and fourth per- 
eiopods simple and much reduced in size; coxal boss on fourth pereiopod 
and projection on coxa of fifth distinctly less prominent. 

First pleopod (Figs. 13, 15) without corneous terminal elements, and 
all more bulbous and less distinctly defined than in holotype. 

Color Notes: Dorsum of carapace pale tan with margins of rostrum 
and postorbital ridges dark brown. Cephalodorsal areas dark brown 
changing to reddish brown in caudal hepatic and gastric areas; ventral 
hepatic region with pair of dark brown oblique bars on cream, tan back- 
ground. Branchiostegites with usual saddle: horns and caudoventral por- 
tion of saddle almost black, bar very narrow dorsally connecting broader 
dorsolateral portions by thin line on extreme caudodorsal margin; area 
below horns pale tan fading to cream. Abdominal terga tan to olive 
green with caudal margins black and each with pair of transverse dor- 
solateral black bars; each of first five abdominal pleura with longitudinal 
black stripe at base and narrow border along margins and each with 
scarlet spot above stripe with cream area between stripe and margin. 
Sixth tergum with caudally emarginate scarlet band cephalically and en- 
tire band caudally. Telson bearing pair of black spots in cephalolateral 



342 Proceedings of the Biological Society of Washington 

corners of caudal section, otherwise tan to dark brown. Outer ramus of 
uropod blackish brown mesially and distally; inner ramus mostly blackish 
brown but with light spot mesially proximal to midlength. Antennules 
and antennae dark brown. Chela brownish black with white to cream- 
tipped tubercles; tips of finger scarlet; carpus and distal portion of merus 
blackish tan dorsally, latter fading proximally through greenish tan to 
cream tan with ischium and coxa mostly cream. Remaining pereiopods 
greenish with brownish markings above, paler below, fading proximally 
to cream tan. 

Type-locality: Tributary to Muckalee Creek (Flint River drainage), 
3.2 miles north of Americus on U.S. Hwy. 19, Sumter County, Georgia. 
This comparatively clear stream, some 10 to 15 feet wide and about two 
feet deep, flows with a moderate current over a sandy and organic bed 
supporting a luxuriant stand of Vallisneria sp. in the unshaded area near 
the bridge. Among the shoreline plants are Salix nigra, Liriodendron tuli- 
pifera, Acer sp., Quercus sp., and Magnolia sp. No other crayfishes were 
collected with P. gibbus in the locality. 

Disposition of types: The holotypic male, form I, the allotypic female, 
and the morphotypic male, form II, are deposited in the Smithsonian In- 
stitution nos. 129804, 129803, and 129805, respectively, as are the para- 
types consisting of 4 ,5 , form I; 7 ^ , form II; 16 $ ; 33 juvenile $ ; and 
41 juvenile $ . 

Size: The largest first form male has a carapace length of 42.2 mm, 
and the smallest, 29.8 mm. The largest specimen available is a second 
form male, the carapace length of which is 46.9 mm. 

Range and Specimens Examined: Procambarus gibbus is known to oc- 
cur in seven localities in the middle Flint River drainage, chiefly in the 
Muckalee Creek watershed, in Georgia. 

CRAWFORD COUNTY— Tributary to Flint River, 1.5 mi. E of Craw- 
ford-Taylor county line, 17 April 1968, Horton H. Hobbs, Jr., coll. (9 $ 
II, 13 2,1 juv. $ , 6 juv. 2 ) . LEE COUNTY— Tributary to Muckalee 
Creek about 7 mi. E of Leesburg, 11 August 1932, E. B. WilHamson, coll. 
( 2 ^ I, 3 ^ II, 2 2 , 4 juv. $ , 1 juv. 2 ) ; Tributary to Muckalee Creek 
about 7 mi. S of Leesburg, 11 August 1932, E. B. WilUamson, coll. (1 $ 
I, 1 5 II, 4 2, 4 juv. $, 1 juv. 2 ) Only data "Lee County," 15 August 
1932, E. B. Williamson, coll. (1 $ I, 1 2); Muckaloochee Creek at 
Smithville, 14 April 1968, Horton H. Hobbs, Jr., coll. (1 ^ II, 2 2, 12 
juv. $ , 14 juv. 2 ) Muckalee Creek, 6 mi. SW of Lee-Sumter County 
line, 15 April 1968, Horton H. Hobbs, Jr., coll. (12). SCHLEY 
COUNTY— Tributary to Muckalee Creek, 3.8 mi. SW of Ellaville on 
State Rte. 153, 15 April 1968, Horton H. Hobbs, Jr., coll. (1 ^ II, 2 2, 
2 juv. $ ). SUMTER COUNTY— Type-locality, 15 April 1968, Horton 
H. Hobbs, Jr., coll. ( 1 ^ I, 2 ^ II, 4 2, 11 juv. $, 25 juv. 2 ). 

Variations: Among the few variations noted, none of which are as- 
sociated with a restricted portion of the range, are occasional cephalo- 



Tivo new crayfish 343 

lateral tubercles on the epistome, telson with one or two spines in caudo- 
lateral corner of cephalic section, inner margin of palm of chela with five 
or six tubercles, upper distal surface of merus of cheliped with two or 
three spines, ischium of cheliped with two to four spines, and hooks on 
ischia of fourth pereiopods in first form male simple or weakly bituber- 
culate. ( See diagnosis for variations in proportions. ) 

Relationships. Procambarus gibbus has its closest relationship with P. 
spiculifer (LeConte, 1856), the range of which completely surrounds 
that of this new species. It shares many features in common with 
LeConte's species but may be readily distinguished from it by the broad, 
distal, sub truncate, caudolateral portion of the first pleopod of the male. 
In P. spiculifer the caudodistal margin of the appendage tapers sharply 
toward the base of the caudal process. 

Life History Notes: First form males have been collected in April and 
August; no females carrying eggs or young have been found. 

Crayfish Associates: Collected with Procambarus gibbus in one or 
more localities were P. paeninsulanus (Faxon, 1914: 369), Faxonella 
clypeata (Hay, 1899: 122), Cambarus d. diogenes Girard, 1852: 88, and 
C. latimanus (Le Conte, 1856: 402). 

Etymology: The name gibbus (L.) meaning protuberant, refers to the 
swollen caudodistal portion of the first pleopod of the male. 

KEY TO THE SPECIES OF THE SPICULIFER GROUP 

The members of the Spiculifer Group of the crayfish genus Procam- 
barus may be recognized readily, for they are the only members of the 
genus occurring the United States which possess two or more cervical 
spines on each side of the carapace. 

First Form Males 
(Based on first pleopods) 

1 Cephalic process well-developed (Figs. 9, 26-34) 2 

1' Cephalic process absent or rudimentary (Figs. 16, 35-38 )__ 12 

2(1) Cephalic process entirely mesial to central projection and 
completely obscured by latter in lateral aspect ( Figs. 32, 

33) P. suttkusi Hobbs, 1953a: 173 

( Choctawhatchee drainage system in Alabama and Flor- 
ida.) 
2' Cephalic process cephalic, lateral, or cephalomesial to 

central projection, never entirely obscured by latter in 

lateral aspect (Fig. 9, 26-31, 34) 3 

3(2') Cephalic process situated cephalomesial to central projec- 
tion (Figs. 28, 35 occasionally) 4 

3' Cephalic process situated cephalic or lateral to central 

projection (Figs. 9, 26, 27, 29-31, 34) 5 



344 Proceedings of the Biological Society of Washington 

4(3) Central projection directed distally (Fig. 28) 

P. penni Hobbs, 1951: 273 

(Pascagoula and Pearl drainage systems in Mississippi 
and Louisiana). 

4' Central projection directed strongly caudodistally (Fig. 

35) P. ouachitae Penn, 1956: 109 

(Arkansas and Ouachita drainage systems in Arkansas). 

5(3') Cephalic process situated lateral to central projection 

(Fig. 31) P. echinatus Hobbs, 1956: 117 

(EdistO' and Salkehatchie drainage systems in South 
Carolina ) . 

5' Cephalic process situated cephalic to central projection 

(Figs. 9, 26, 27, 29, 30, 34) 6 

6(5') Cephalic process almost straight, directed distally or ceph- 

alodistally (Figs. 9, 26-28) 7 

6' Cephalic process recurved, often hooding central projection 

(Figs. 29, 30, 34) 10 

7(6) Mesial process not extending distally beyond tip of central 

projection (Fig. 28) P. penni [see couplet 4] 

7' Mesial process extending distally beyond tip of central pro- 
jection (Figs. 9, 26, 27) 8 

8(7') Caudal knob truncate in caudal aspect, obscuring caudal 

process (Figs. 5, 9) P. elegans, new species 

(Tributaries of the Ouachita River in northern Louis- 
iana ) . 

8' Caudal knob irregular in caudal aspect with caudal process 

evident (Figs. 26, 27) 9 

9(8') Central projection recurved, accessory process small (Fig. 

26) P. ablusus Penn 1963: 121 

(Hatchie drainage systems in Mississippi and Tennes- 
see). 

9' Central projection almost straight, accessory process con- 
spicuous (Fig. 27) P. natchitochae Penn, 1953: 5 

(Bayou Teche, Red, and Calcasieu drainage systems in 
Arkansas, Louisiana, and Texas). 
10(6') Caudal knob distinct from caudal process and knoblike 

(Fig. 29) P. vioscai Penn, 1946: 27 

( From the Red drainage systems in Arkansas and Louis- 
iana eastward to, but not in, the Pearl River system in 
Mississippi ) . 
10' Caudal knob not clearly distinct from caudal process and 

subacute (Figs. 30, 34) 11 

11(10') Distal portion of pleopod tapering conspicuously in lateral 
aspect, mesial process directed caudodistally (Fig. 34) 
P. versutus (Hagen, 1870: 51) 



Two new crayfish 345 

11' Distal portion of pleopod tapering only slightly in lateral 

aspect, mesial process directed caudally at angle greater 

than 45° to shaft of appendage (Fig. 30) 

P. dupratzi Penn, 1953: 1 

( Trinity, Red, Neches, Sabine, and Calcasieu drainage 
systems in Arkansas, Texas, and Louisiana ) . 

12(1') Central projection arising from level distinctly proximal to 

base of caudal process (Figs. 16, 38) 13 

12' Central projection not arising from level distinctly proxi- 
mal to base of caudal process (Figs. 35-37) 14 

13(12) Caudodistal portion of shaft subtruncate with caudal ele- 
ment and central projection situated on cephalic half of 

tip (Fig. 16) P. gibbus new species 

(Tributaries to the Flint River in southwestern Geor- 
gia). 

13' Caudodistal portion of shaft tapering with caudal element 

and central projection constituting almost entire tip 

(Fig. 38) P. spiculifer (LeConte, 1856): 401 

(From the Alabama drainage system eastward to the 
Savannah system but not south of the Suwannee drain- 
age in Florida). 

14(12') Distolateral surface of shaft with longitudinal excavation 
extending proximally from central projection (Fig. 36) 

P. lagniappe Black, 1968: 5 

(Sucarnooche Creek drainage — to Tombigbee River — in 
Kemper County, Mississippi and Sumter County, Ala- 
bama ) . 

14' Distolateral surface of shaft without longitudinal excava- 
tion (Figs. 35, 37) 15 

15(14') Caudal knob conspicuous and caudal process large (Fig. 

35) P. ouachitae Penn [see couplet 4] 

15' Caudal knob vestigial and caudal process not large (Fig. 

37) P. mneyi Hobbs, 1953b: 412 

(Upper Savannah River drainage in Georgia and South 
Carolina and headwaters of the Ocmulgee River in De- 
Kalb County, Georgia). 

Females 

( For bibliographic citations and ranges, see Key to Males ) 
1 Cephalic portion of annulus ventralis obscured by promi- 

nent multituberculate lobes of sternum immediately 

cephalic to it 2 

1' Cephalic portion of annulus ventralis not obscured by 
prominent multituberculate lobes of sternum immediately 
cephalic to it although single tubercles may project 
slightly over its margin 7 



346 Proceedings of the Biological Society of Washington 

2(1) Sternum cephalic to annulus ventralis with at least one 
subdigitiform tubercle extending caudomesially beyond 
transverse ridge of annulus P. suttkusi Hobbs 

2' Sternum cephalic to annulus ventralis without digitiform 

tubercles 3 

3(2') Annulus without or with very shallow median longtudinal 

furrow cephalically P. lagniappe Black 

3' Annulus ventralis with conspicuous median longitudinal 

furrow cephalically 4 

4(3') Median longitudinal furrow of annulus flanked by oblique 

ridges 5 

4' Median longitudinal furrow of annulus flanked by trans- 
verse ridges 6 

5(4) Sinus of annulus with at least one acute angular bend .— 

P. dupratzi Penn 

5' Sinus of annulus with no acute bends P. ouachitae Penn 

6(4') Tubercular lobes extending caudally from sternum ce- 
phalic to annulus almost contiguous from base to end; 
carapace usually with more than two cervical spines 
P. echinatus Hobbs 

6' Tubercular lobes extending caudally from sternum cephalic 

to annulus each forming broad convex arc from median 

line; carapace with only two cervical spines 

P. raneyi Hobbs 

7( 1') Sternum cephalic to annulus devoid of tubercles or lobes — . 8 

7' Sternum cephalic to annulus always with tubercles or small 

lobes 9 

8(7) Areola at least five times longer than broad 

P. elegans new species 

8' Areola less than five times longer than broad 

P. ablusus Penn 

9(7') Cephalolateral portions of annulus with pair of narrow 

ridges paralleling cephalolateral margins _— P. vioscai Penn 

9' Cephalolateral portions of annulus never with pair of nar- 
row ridges paralleling cephalolateral margins 10 

10(9') Caudal margin of annulus transverse P. penni Hobbs 

10' Caudal margin of annulus subtriangular, or with caudo- 

median prominence 11 

11(10') Caudal margin of annulus forming gentle arc 

P. natchitochae Penn 

11' Caudal margin of annulus broadly subtriangular or- with 

distinct caudomedian prominence 12 

11(10') Median longitudinal furrow of annulus deep cephalically 

and with very high lateral walls P. gibbus, new species 

11' Median longitudinal furrow of annulus shallow cephalically 

and with walls not conspicuously high ..P. spiculifer LeConte 



Two new crayfish 347 

Discussion: This group, comprising 14 species of stream-dwellers, oc- 
cupies a range in the Coastal Plain and Piedmont provinces extending 
from the Trinity River, Texas, eastward to the Edisto River, South Caro- 
lina, and Suwannee River, Florida. Although occasionally encountered in 
areas of streams where there is no obvious current, they are not known to 
invade lentic habitats. 

The ranges of five species of the group — P. ablusus, P. echinatus, P. 
lagniappe, P. ouachitae and P. penni, — appear to be allopatric; whereas, 
those of the remaining species overlap that of one or more members of 
the group. The range of P. vioscai is partially shared by P. dupratzi, P. 
natchitochae, and P. elegans, while that of P. spiculifer includes parts of, 
or surrounds, that of P. gibbus, P. raneyi, P. suttkusi, and P. versutus. De- 
spite the overlapping ranges^ only two combinations of these species have 
been collected in a single locality — P. elegans and P. vioscai in Louisiana, 
and P. spiculifer and P. versutus in Alabama and Florida. 

Acknowledgments: I wish to thank Lt. T. H. Nickerson for presenting 
me with the first specimen of Procambarus elegans I had seen, and I am 
indebted to Drs. N. H. Douglas, W. J. Harman, L. W. Lowe, and W. R. 
Taylor, for assistance in obtaining additional representatives of this spec- 
ies. I also wish to thank Drs. Fenner A. Chace and J. F. Fitzpatrick for a 
number of valuable suggestions and criticisms. For the exceptionally fine 
illustrations of this species (Figs. 1-4), I extend my genuine appreciation 
to Mrs. Carolyn B. Cast. 

LiTERATXJRE CiTED 

Black, Joe B. 1968. A new crawfish of the genus Procambarus from 
Mississippi (Decapoda, Astacidae). Tulane Studies in Zool. 
and Botany, 15( 1 ) : 5-9, 12 figs. 

Faxon, Walter. 1898. Observations on the Astacidae in the United 
States National Museum and in the Museum of Comparative 
Zoology, with descriptions of new species. Proc. U.S. Nat. 
Mus., 20 (1136): 643-694, pis. LXII-LXX. 

. 1914. Notes on the crayfishes in the United States National 

Museum and the Museum of Comparative Zoology with de- 
scriptions of new species and subspecies to which is appended 
a catalogue of the known species and subspecies. Mem. Mus. 
Comp. Zool., Harvard Coll., 40(8): 351-427, 13 pis. 

GiRARD, Charles. 1852. A revision of the North American Astaci, with 
observations on their habits and geographical distribution. 
Proc. Acad. Nat. Sci., Philadelphia, 6: 87-91. 

Hagen, Hermann A. 1870. Monograph of the North American Astacidae. 
lUus. Cat. Mus. Comp. Zool., Harvard Coll., (3): 1-109, 11 
pis. 

Hay, William Perry. 1899. Description of two new species of crayfish. 
Proc. U.S. Nat. Mus., 22 (1187): 121-123, 2 figs. 



348 Proceedings of the Biological Society of Washington 

HoBBS, HoRTON H,, Jr. 1945. Two new species of crayfishes of the genus 
Cambarellus from the Gulf Coastal states, with a key to the 
species of the genus (Decapoda, Astacidae). Amer. Midi. 
Nat., 34(2): 466-474, 26 figs. 

. 1948. A new crayfish of the genus Camharus from Texas, 

with notes on the distribution of Camharus fodiens (Cottle). 
Proc. U.S. Nat. Mus., 98 (3230): 223-231, fig. 17. 

. 1951. A new crayfish of the genus Procambarus from Loui- 
siana with a key to the species of the Spiculifer Group. Journ. 
Wash. Acad. Sci., 41(8): 272-276, 11 figs. 

. 1953a. A new crayfish of the genus Procambarus from Al- 
abama and Florida (Decapoda, Astacidae). Proc. Biol. Soc. 
Wash., 66: 173-178, 10 figs. 

. 1953b. On the ranges of certain crayfishes of the Spiculifer 

Group of the genus Procambarus, with the description of a 
new species (Decapoda, Astacidae). Journ. Wash. Acad. 
Sci., 43(12): 412-417, 13 figs. 

. 1956. A new crayfish of the genus Procambarus from South 

Carolina (Decapoda, Astacidae). Journ. Wash. Acad. Sci., 
46(4): 117-121, 17 figs. 

. 1958. The evolutionary history of the Pictus Group of the 

crayfish genus Procambarus (Decapoda, Astacidae). Quart. 
Journ. Fla. Acad. Sci., 21( 1 ) : 71-91, 20 figs. 

LeConte, John. 1856. Descriptions of new species of Astacus from 
Georgia. Proc. Acad. Nat. Sci., Philadelphia, 7: 400-402. 

Penn, George Henry, Jr. 1946. A new crawfish of the genus Pro- 
cambarus from Louisiana (Decapoda: Astacidae). Journ. 
Wash. Acad. Sci., 36( 1 ) : 27-29, 6 figs. 

. 1953. Two new crawfishes of the genus Procambarus from 

Texas, Louisiana, and Arkansas (Decapoda, Astacidae). 
Amer. Mus. Novitates (1636): 1-10, 19 figs. 

. 1956. A new crawfish of the genus Procambarus from Ar- 
kansas (Crustacea, Astacidae). Lloydia, 19(2): 109-119, 17 
figs. 

. 1963. A new crawfish from the Hatchie River in Mississippi 

and Tennessee (Decapoda, Astacidae). Proc. Biol. Soc. 
Wash., 76: 121-126, 10 figs. 



B4X 

^^ i54 3 October 1969 

BIOLOGICAL SOCIETY OF WASHINGTON 20 V969 



CS-BI 



A NEW SPECIES OF MEIACANTHUS ( PISCES :^LEN- 
NIIDAE: NEMOPHIDINAE) FROM THE RED SEA, 
WITH A REVIEW OF THE INDIAN OCEAN SPECIES 

By William F. Smith- Vaniz 
Institute of Marine Sciences, University of Miami 

Two of the three species of Meiacanthus described since the 
genus was first recognized (Norman, 1943) have come from 
the Indian Ocean, an area from which the genus was previously 
unknown. The purpose of this paper is to review the Indian 
Ocean species and describe an additional new species known 
only from the Red Sea. Information on the other members 
of the genus will be published upon completion of a study now 
in progress treating the subfamily to which Meiacanthus be- 
longs. The description of the new Meiacanthus appears at this 
time so that the name will be available to other workers, par- 
ticularly those interested in Red Sea fauna. 

Methods of counting and measuring follow those of Springer 
(1967). The caudal ratio was obtained by measuring the 
length of the shortest, undamaged, middle caudal ray and 
arithmetically dividing this measurement into the length of the 
longest, undamaged outer caudal ray. Specimens with apparent 
injuries to both lobes of the caudal fin were not measured. All 
counts of median fin elements were obtained from radiographs. 
Measurements were taken with needle-point dial calipers to the 
nearest tenth of a millimeter. 

For the loan of specimens in their care I thank Dr. Reeve M. 
Bailey, Museum of Zoology, University of Michigan (UMMZ), 
Mrs. Margaret M. Smith, Rhodes University (RU), and Dr. 
Victor G. Springer, Smithsonian Institution (SI). Drs. C. 
Richard Robins and Victor G. Springer critically read the manu- 
script. 

Contribution No. 1094 from the Institute of Marine Sciences, University of Miami. 

25— Proc. Biol. Soc. Wash., Vol. 82, 1969 (349) 



350 Proceedings of the Biological Society of Washington 



S 
2 



New Red Sea blenny 351 

Meiacanthus nigrolineatus new species 
Fig. 1 

Holotype: USNM 200301. Female 31.7 mm standard length, from the 
Red Sea, Straits of Jubal (27°14'34"N., 33°53'55"E.), depth 0-8 ft., 3 
January 1965, H. A. Fehlmann (sta. HA-31). 

Paratypes: USNM 200601. (4, 30.4-34.0, including 1 cleared and 
stained), data same as for holotype. USNM 201480. (3, 36.0^53.0), Red 
Sea (27°17'32"N., 33°48'52"E.), depth 0-20 ft., 7 January 1965, H. A. 
Fehlmann (sta. HA-36). USNM 201485. (1, 41.3), Red Sea (27° 18' 
50"N., 33°47'35"E.), depth 0-15 ft., 10 January 1965, H. A. Fehlmann 
(sta. HA-38). 

Description: A medium-sized species of Meiacanthus with outermost 
rays of caudal fin moderately elongated; caudal ratio 1.5-2.0 in males, 
1.3-1.5 in females. Middle ray of each pelvic fin only slightly longer than 
adjacent rays in either sex; pelvic length 10.9^13.1, average 12.1 in 
standard length. A single pore at mid-dorsal point of supratemporal sen- 
sory canal. Pores of preopercular series without cirri. Lateral line con- 
sisting of four or five short, separate, longitudinally bipored tubes that end 
beneath base of second or third segmented dorsal ray. 

Dorsal rays IV, 24 or 25; segmented anal rays 14 or 15; total pectoral 
rays 26 to 28, usually 26 or 27. Vertebrae 12 -f 21 or 22 = 33 or 34, 
usually 34; epipleural ribs on vertebrae 1 through 14 or 15; pleural ribs 
on vertebrae 3 through 12. Premaxillary and dentary incisors 17 to 21, 
the incisors increasing in number with increase in size as is typical for all 
members of the genus. Gill-rakers 7 or 8; pseudobranchial filaments 
6 to 8. 

Pigmentation: The pigmentation varies little (Fig. 1). Sexual di- 
chromatism was not apparent in the type series of specimens. Diagnostic 
features of the color pattern are given in Table 1. 

Relationship : Since a revision of the genus Meiacanthus is now in prog- 
ress, a discussion of relationships will not be given at this time. Com- 
parison of nigrolineatus with the two species of the genus known from the 
Indian Ocean is given in Tables 1-2. 

Etymology: The name nigrolineatus refers to the narrow black stripe 
which is the most distinctive character of the species. 

Discussion: M. nigrolineatus is known only from the Red Sea and will 
probably prove to be endemic. Marshall ( 1952 ) has commented on the 
high incidence of endemism in the Red Sea. He noted that a number of 
Red Sea endemics differed from closely related Indian Ocean forms in 
typically having lower meristic counts and more slender bodies. Meiacan^ 
thus nigrolineatus conforms well to this pattern of differentiation (see 
Table 2). 

THE STATUS OF INDIAN OCEAN SPECIES OF MEIACANTHUS 

The first species of Meiacanthus described from the Indian Ocean was 

M. mossamhicus (Smith, 1959). In the original description it was stated 






352 Proceedings of the Biological Society of Washington 






a-^^^ 



a s 



X .5 "" o 



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New Red Sea blenny 



353 



Table 2. Distribution of fin ray counts of the three species of Meia- 
canthus from the Red Sea and Indian Ocean.^ 





Dorsal spines 


Segmented dorsal rays 
24 25 26 27 


Total dorsal rays 




4 5 


28 


29 30 31 


nigrolineatus 
mossambicus 
smithi 


(9) 

3 13 
9 


2 (7) 

1 10 4 1 
1 8 


2 


(7) 

2 10 4 
1 8 






Segmented 
anal rays 


Total dorsal and anal 
spines and rays 




Total pectoral rays 




14 15 16 17 


44 45 46 47 48 49 


26 27 28 29 30 


nigrolineatus 
mossambicus 
smithi 


2 (7) 
6 10 
1 8 


1 2 (6) 

16 5 4 
1 8 




(5) 2 2 

15 1 
16 2 



1 Counts for holotype of M. nigrolineatus in parentheses. 

to be "closely related or possibly identical" with M. atrodorsalis (Giin- 
ther), a widely distributed Indo-Pacific species. The color pattern of 
mossambicus will readily distinguish it from all known species of the ge- 
nus. The typical dorsal spine count of 5 will distinguish it from M. atro- 
dofsalis, and all but one other member of the genus. 

A second species of Meiacanthus from the Indian Ocean was described 
by Klausewitz ( 1962) on the basis of a single specimen from the Mal- 
dive Islands. This species which he named sm,ithi was not compared to 
Indo-Pacific members of the genus. In most features of pigmentation 
smithi is remarkably similar to M. atrodorsalis and their meristic charac- 
ters broadly overlap. The only pigment character that appears to separate 
all specimens of either species is the presence of a conspicuous dark spot 
( slightly smaller than diameter of pupil ) in the axil of the pectoral fin in 
atrodorsalis. This spot which is clearly evident in the 67 specimens ( 19.8- 
67.2 mm SL) of atrodorsalis that I have examined does not occur in 
smithi. 

The two species can also be distinguished from each other on the basis 
of several morphometric characters associated with sexual dimorphism. 
In M. atrodorsalis the pelvic-fin rays exhibit no sexual dimorphism, the 
middle segmented ray being similarly reduced in both sexes, while in 
smithi the middle ray is greatly elongated in males ( see Table 1 ) . In con- 
trast to smithi the caudal fin exhibits sexual dimorphism in atrodorsalis. 
Furthermore, although the caudal ratio is smaller in females of atrodor- 
salis than in the males, it is larger than in either sex of smithi. The caudal 
ratios for a number of specimens of atrodorsalis is as follows: In 26 
males (38.8-67.2 SL) the caudal ratio was 2.12-4.19, average 2.84; in 
17 females (36.0-63.7 SL) the caudal ratio was 1.66-3.05, average 2.22; 
in 17 specimens of either sex (19.8-34.5 SL) the caudal ratio was 1.62- 
2.46, average 1.96. In smithi the outer caudal rays are not elongated in 
either sex and the caudal ratio is 1.0. 



354 Proceedings of the Biological Society of Washington 

Specimens Examined: Meiacanthus mossamhicus. Mombasa: RU un- 
cat., 25 Feb. 1964 (1, 42.8); Zanzibar: RU uncat., sta. Z. 576 (1, 28.7); 
Mozambique: RU uncat., 28 Aug. 1950 (2, 66.9-68.9); Pinda: RU un- 
cat. sta. B.P. 1793 (1, 58.3), RU uncat. sta. B.P. 1192 (1, 62.1), RU 
uncat. sta. B.P. 1112 (1, 62.5), RU uncat. sta. B.P. 319 (1, 48.5); prob- 
ably Pinda: RU uncat. (1,55.5); Wamizi Is.; RU uncat. (1,51.6); Comoro 
Is.: Anjouan Is.: USNM 210483 (1, 69.2); Pamanzi Is.: USNM 201484 
(2,50.0-53.3); Madagascar: East Ambariobe, Nossi-Be': USNM 201493 
(1, 37.8); Tulear UMMZ 185821 (1, 60.3), UMMZ 185968 (1, 54.1). 

Meiacanthus smithi. Thailand: Phuket, Patong USNM 201489 (1, 
37.9); Indonesia: Mentawei Is. USNM 199483 (2, 56.1-64.4); Malacca 
Straits USNM 199422 (6,32.3-50.2). 

Literature Cited 

Klausewitz, Wolfgang. 1962. Meiacanthus smithi n. sp. aus dem In- 
dischen Ozean. Senck. Biol., 43(1): 17-19, 1 fig. 

Marshall, Norman B. 1952. The 'Manihine' Expedition to the Culf of 
Aquaba 1948-1949 IX. Fishes. Bull. Brit. Mus. Nat. Hist., 
1 (8): 221-252, 3 figs. 

Norman, John R. 1943. Notes on the Blennioid fishes, I: A provisional 
synopsis of the genera of the family Blenniidae. Ann. Mag. 
Nat. Hist., 10(11): 793-812. 

Smith, John L. B. 1959. Fishes of the families Blenniidae and Salari- 
idae of the western Indian Ocean. Rhodes Univ. Ichth. Bull., 
14: 229-252, pis. 14-18. 

Springer, Victor G. 1967. Revision of the circumtropical shorefish ge- 
nus Entomacrodus (Blenniidae: Salariinae). Proc. U.S. Nat. 
Mus., 122(3582): 1-150, 30 pis. 



'^^ 



QH 
1 

B4X 
NH 



U 7 2 



55-372 3 October 1969 



PROCEEDINGS 
OF THE 

BIOLOGICAL SOCIETY OF WASHINGTON 



CITHARICHTHYS ABBOTTI, A NEW FLATFISH 

(BOTHIDAE) FROM THE SOUTHWESTERN GULF OF 

MEXICO 

By C. E. Dawson 

Gulf Coast Research Laboratory, 

Ocean Springs, Mississippi 

I have recently collected a number of specimens of a flatfish, 
referable to the bothid genus Citharichthys Bleeker, from 
coastal waters of Veracruz and Campeche, Mexico. It was im- 
possible to identify this fish by means of existing revisionary 
studies or keys (Parr, 1931; Norman, 1934; Gutherz, 1967) and 
comparison with its most closely related western Atlantic con- 
geners reveals significant differences in a number of meristic 
and morphological characters. In view of its apparent abun- 
dance on the Mexican coast and the possibility of its confusion 
with related Gulf of Mexico and Caribbean forms, I take this 
opportunity to describe this new species and to provide com- 
parative notes on C. spilopterus Giinther and C. arenaceus 
Evermann and Marsh. 

Type material has been deposited in the collections of the 
following institutions: United States National Museum (US- 
NM); Academy of Natural Sciences of Philadelphia (ANSP); 
Museum of Comparative Zoology ( MCZ ) ; University of Miami, 
Institute of Marine Science (UMML); Instituto Nacional de 
Investigaciones Biologico Pesqueras, Mexico (INIBP) and 
the Gulf Coast Research Laboratory ( GCRL ) . 

Measurements were made with dial calipers or ocular microm- 
eter and recorded to the nearest 0.1 mm. Morphometric 
criteria requiring special definition are as follows: standard 
length (SL) — anteriormost tip of premaxillary to rear of hy- 
pural; head length (HL) — tip of premaxillary to posteriormost 
fleshly margin of opercle; body depth — ^maximum vertical dis- 

26— Proc. Biol. Soc. Wash., Vol. 82, 1969 (355) 




356 Proceedings of the Biological Society of Washington 

tance between dorsal and anal fin bases; pectoral fin length — 
base of first ( dorsad ) ray to tip of longest ray; pelvic fin length 
— insertion of anteriormost ray to tip of longest ray; diameter of 
orbit — maximum longitudinal distance between bony margins 
of right ( dorsad ) orbit; eye diameter — maximum longitudinal 
diameter of left eye; snout length — anterior margin of orbit to 
tip of premaxillary; postorbital length — posterior margin of 
orbit to tip of opercle; upper jaw length — tip of premaxillary to 
posteroventral margin of maxillary; lower jaw length — antero- 
dorsal tip of dentary to rear of retroarticular angle; length of 
1st dorsal ray — from anterior angle of insertion to tip. Scale 
counts include the number of pored lateral line scales from 
upper opercular angle to rear of hypural. Gill raker counts are 
those of the 1st left side arch and include rudiments as well as 
fully developed rakers. A raker located at the angle is included 
in the count of the lower arch. Tabulated meristic counts are 
based on material examined during this study. Description and 
drawings of dentition are from cleared, stained and dissected 
specimens (100-115 mm SL). Statistical analyses were run on 
an IBM 1130-2B 8K computer. 

Citharichthys abbotti new species 
(Fig. 1) 

Holotype: USNM 203735; 142.6 mm SL; Mexico, Veracruz, from eddy 
current at mouth of Laguna de Zontecomapan; ichthyocide collection 
over sand bottom in depths up to 2 meters; 1 September 1967; C. E. 
Dawson and W. Abbott coll. 

Paratypes: ANSP 112317; (2) 74-83 mm SL. GCRL 2436; (3) 64- 
132 mm SL. GCRL 2808; (2) 103-112 mm SL, cleared and stained. 
MCZ 46368; 83 mm SL. UMML 24484; (2) 79-90 mm SL. USNM 
203736; (6) 28-127 mm SL. INIBP uncat.; 76 mm SL. Collection data 
of foregoing paratypes as for holotype. USNM 203737; (12) 24-65 mm 
SL; from tidepool on north side of south jetty at Alvarado, approx. 
18°47'30"N, 95°45'07"W; 7 June 1968; C. E. Dawson coll. USNM 
203738; (6) 23-94 mm SL; Campeche, east shore of Rio San Pedro, near 
mouth, approx. 18°41'45"N, 92°27'15"W; 10 June 1968; C. E. Dawson 
coll. 

Other material: Mexico, Veracruz— CCRL 2382; (5) 13-17 mm SL; 
Isla da Amor, Boca del Rio; 4 September 1967; C. E. Dawson coll. GCRL 
2368, 2685; (3) 126-138 mm SL; Gulf of Mexico beach just south of 
Boca del Rio; 4 September 1967; C. E. Dawson and seine crew coll. 
GCRL 2435; (5) 13-18 mm SL; Rio Jamapa, approx. 0.5 mi. from mouth; 



New Gulf of Mexico flatfish 



357 




Fig. 1. Citharichthys ahhotti USNM 203736; paratype; 116.0 mm SL. 



4 September 1967; C. E. Dawson and S. Romay coll. GCRL 2733; (20) 
15-75 mm SL; Isla da Amor, Boca del Rio, approx. 19°05'32"N, 96°06' 
W; 5 June 1968; C. E. Dawson coll. GCRL 2759; (27) 20^62 mm SL; 
north shore of Isla Pajaritos, Rio Coatzacoalcos, approx. 18°07'21"N, 
94°25'02"W; 8 June 1968; C. E. Dawson coll. GCRL 2758, 2807; (45) 
16-45 mm SL, (3) cleared and stained; mouth of Laguna de Zontecom- 
apan; 16 August 1968; C. E. Dawson coll. 

Mexico, Campeche — GCRL 2802; 51 mm SL; Zacatal, just north of 
ferry landing, approx. 18°37'N, 91°51'45"W; 9 June 1968; C. E. Dawson 
coll. 

Diagnosis: A species of Citharichthys lacking osseous protuberances or 
spines on snout or orbital rims; modally with 22 caudal vertebrae and 
combinations of 76 dorsal and 56 anal fin rays, 20 gill rakers on the first 
arch and 44-45 lateral line scales; without prominent or distinctive colora- 
tion; first dorsal ray inserted close to anterior blind side naris, its length 
averages 8 percent of SL; snout about 30 percent longer than eye, sub- 
equal to diameter of orbit; body depth averages 47 percent of SL, left 
pelvic fin averages 12 percent of SL and the upper jaw averages 37 per- 
cent of HL. 

Description: Dorsal rays 72^81, usually 75-78; anal rays 52-60, usually 
55-56, gill rakers 5^8 + 13-16 = 18-23, usually 6 + 14 = 20; left pec- 
toral fin 9-11, usually 10; right pectoral fin 8-10, usually 9; pelvic fins 
6; segmented caudal fin rays 17; lateral line scales 42-49, usually 44-45; 
vertebrae 9-11 + 21-23 = 31-33, usually 10 + 22 = 32. 

Measurements (mm) of holotype are as follows: caudal fin length 
32.6, depth of caudal peduncle 19.3, body depth 69.2, left pectoral fin 
length 18.1, right pectoral fin length 16.7, left pelvic fin length 18.1, right 






358 Proceedings of the Biological Society of Washington 



Table 1. Frequency distribution of dorsal and anal fin rays in Cith- 

arichthys abbotti, C. arenaceus and C. spilopterus. (Holotype marked *; 

X = mean; a = standard deviation). 





N 




Dorsal rays 




X 




Species 


70 


71 72 73 74 75 76 77 78 79 80 81 


82 


(T 


abbotti 

arenaceus 

spilopterus 


132 

45 
87 


1 


1 5 13 23 30 23 23* 11 2 1 
2 11 10 9 7* 3 2 

2 1 9 20 20 16 17* 6 


3 


76.3 
73.4 
78.3 


1.73 
1.60 
1.78 






N 




Anal rays 




X 




Species 


51 


52 53 54 55 56 57 58 59 60 61 62 




(T 


abbotti 

arenaceus 

spilopterus 


132 
46 

87 


1 


1 5 13 37 35 22 14* 4 1 
11 8 14 9* 3 

8 9 20 23 17* 7 3 




55.8 
53.6 

58.7 


1.46 
1.29 
1.50 



pelvic fin length 16.7, length of first dorsal ray 11.1, head length 40.8, 
diameter of bony orbit 7.7, eye diameter 5.8, width of bony interorbital 
2.1, anterodorsal margin of orbit to dorsal fin base 3.0, postorbital 
length 26.2, upper jaw length 15.5, lower jaw length 18.1. See Tables 1-4 
for additional counts and measurements. 

Body elongate, oval, its greatest depth (44-51 percent of SL) near a 
vertical from 10th or 11th anal ray; slender, its greatest thickness 7.4 per- 



Table 2. Frequency distribution of upper, lower, and total gill rakers 

in C. abbotti, C. arenaceus, and C. spilopterus. (Holotype marked *; 

X = mean; a = standard deviation). 







N 








Upper arch 








X 




Species 


3 


4 


5 6 


7 


8 


a 


abbotti 

arenaceus 

spilopterus 




89 
40 
85 




1 
19 


6* 

53* 


10 44* 
19 12 
13 


32 

2 


3 




6.3 
5.2 
3.9 


0.72 
0.85 
0.61 








N 








Lower arch 








X 




Species 


11 


12 


13 14 


15 


16 


(T 


abbotti 

arenaceus 

spilopterus 




89 
40 
85 




5 


9 
19 


18 33* 
23* 7 
37 14* 


30 
1 
10 


8 




14.3 
13.0 
13.0 


0.90 
0.72 
1.05 






N 










Total 












Species 


13 


14 


15 


16 


17 18 19 


20 


21 


22 


23 


X (T 


abbotti 

arenaceus 

spilopterus 


89 
40 
85 


1 


4 


1 
12 


3 
19 


1 12 
10* 16 6 
27 16* 6 


31* 
3 


27 

1 


13 


5 


20.6 1.10 
17.9 1.22 
16.6 1.32 



New Gulf of Mexico flatfish 359 

Table 3. Frequency distribution of lateral line scales in Citharichthys 

ahhotti, C. arenaceus, and C. spilopterus. (Holotype marked *; x = 

mean; <t r= standard deviation). 





N 








Lateral line scales 








X 




Species 


42 


43 


44 


45 


46 


47 


48 


49 


50 


cr 


abbotti 


51 


1 


6 


14* 


14 


7 


7 


1 


1 




44.9 


1.46 


arenaceus 


27 








1 




7 


7* 


6 


6 


48.2 


1.30 


spilopterus 


28 




2 


8 


11 


3 


2 


2 






45.0 


1.29 



cent of SL; caudal peduncle short, its depth averages 13 percent of SL. 
Anterodorsal profile broadly rounded, a slight notch before the eye; 
lower jaw slightly included anteriad but with the anteroventral angle of 
dentary projecting slightly in advance of premaxillary; posterior edge of 
maxillary reaches a vertical through the posterior 1/6 of left eye and 
posterior edge of right eye; right eye slightly in advance of left, well re- 
moved from dorsal fin base; fleshy interorbital broad, its width somewhat 
more than 1/2 eye diameter; bony interorbital with a slight ridge along 
margin of left orbit, not continued on snout which is broad and distinctly 
longer than eye. Dorsal fin originates on right side (Fig. 2), about 2/3 of 
eye diameter in advance of the right eye, just above or slightly in ad- 
vance of the anterior naris and about 1/2 an eye diameter ventrad of the 
snout margin; first two dorsal rays closely spaced, interspace between 1st 
and 2nd rays about 1/2 that between 2nd and 3rd; adpressed 1st dorsal 
ray reaches base of 7th ray; longest dorsal and anal rays about 12 and 13 
percent of SL, respectively; left side nares small (0.6 mm in diameter), 
the posterior with a slightly raised margin, the anterior a short tubule 
with a flaplike expansion of its rear margin and a slender tendril ante- 
riad; right side nares similar to those of left but the diameter of the 
posterior naris is slightly larger (0.7 mm) than that of the anterior (0.5 
mm). Left pectoral fin obliquely rounded, the 4th and 5th rays the 
longest, reaches a vertical from 32nd or 33rd dorsal ray; right pectoral 
fin smaller, narrowly rounded, the middle rays the longest; left pelvic 
fin broad, its 2nd and longest ray reaches to between bases of 5th and 
6th anal rays; right pelvic fin smaller; its 4th ray reaches base of 2nd anal 
ray; anus and urinary papilla on right side adjacent to anal fin origin. 

Table 4. Frequency distribution of caudal vertebrae in Citharichthys 
abbotti, C. arenaceus, and C. spilopterus. All data are from radiographs 
and include terminal vertebra bearing hypural. ( Holotype marked * ) . 





N 




Caudal vertebrae 






Species 


21 


22 


23 


24 


25 


X 


abbotti 


124 


19 


96* 


9 






21.9 


arenaceus 


49 


3 


38* 


8 






22.1 


spilopterus 


68 






11 


54 


3 


23.8 



I 



360 Proceedings of the Biological Society of Washington 






New Gulf of Mexico flatfish 361 

Gill rakers (Fig. 2) somewhat crowded, of moderate length, variable, 
but usually slender, those of upper arch generally well-developed, oc- 
casionally with a few serrations on dorsal margins of one or two lower 
arch rakers. 

Scales adherent; about 17 above and 23 below lateral line at its inflec- 
tion; right side scales cycloid; primary left side squamation predominately 
ctenoid, occasionally cycloid on posterior body and caudal peduncle; 
cteni of moderate length, best developed on head and anterior body; 
secondary squamation apparently cycloid throughout. Snout, orbits and 
anterior 2/3 of interorbital naked; scales continue dorsad above and 
slightly in advance of right eye; largest specimens with 5 or 6 scales on 
posterior expansion of maxillary; modified scales continue to a greater 
or lesser extent along all fin rays; pectoral fin base apparently naked; 
left side squamation complete over remainder of body. Lateral line 
slightly elevated above pectoral fin, distinct, usually with two or three 
secondary scales between each pored scale (Fig. 2); secondary squama- 
tion sparse or absent elsewhere, usually present only in larger fish, re- 
stricted to a single small cycloid scale between lateral angles of adjacent 
primary scales, apparently most frequent on posterior body dorsad of 
lateral line. 

Upper and lower jaws with a single outer series of fixed teeth and one 
or more inner series of movable teeth, oriented at approximately 90° 
from the fixed series, and located on the intraoral integument; each 
series somewhat laterally offset from the next (Figs. 3 and 4). Movable 
teeth usually in two series, occasionally three in front; teeth of the 1st 
series, located immediately behind the fixed teeth, are slightly smaller 
and straighter than their fixed counterparts; teeth of the innermost series 
the smallest and straightest. With about 40 close-set, sharp, recurved 
fixed conical teeth on left side of upper jaw; the anteriormost 4-5 slightly 
enlarged. Right side of upper jaw with about 44 fixed teeth, similar to 
but somewhat more slender than those of the left side; the 5-6 enlarged 
anterior teeth are about 1/5 longer than those of the left. Left side of 
lower jaw with 15-19 somewhat separated, recurved conical teeth in the 
fixed series; the first three the largest. Right side of lower jaw with 27-29 
teeth in the outer series; the anteriormost 6-7 much the larger; denti- 
tion continued much further posteriad than on the left dentary. Teeth of 
all series gradually decrease in size posteriad. 



Fig. 2. Upper: Dorsal fin origin, relative lengths of 1st dorsal fin rays, 
together with insert details of right side nares in Cithar- 
ichthys abbotti (left) and C. spilopterus (right). Middle: 
Usual configuration of gill rakers on 1st left gill arch of C. 
abbotti. Lower: Semidiagrammatic delineation of primary 
lateral line scales and associated secondary squamation in 
C. abbotti. 



362 Proceedings of the Biological Society of Washington 






New Gulf of Mexico flatfish 363 

Coloration of recently preserved alcoholic specimens similar to that in 
life. Left side ground color a light sandy-tan; densely speckled with 
small brown blotches; scales frequently margined with a fine, very dark 
brown line; fin rays lined with small brown blotches, most specimens with 
a moderately distinct, darker, blotch on basal one-third of every 5th-8th 
dorsal and anal fin ray; interradial membranes usually pale; a small, 
sometimes indistinct, dark brown spot at upper opercular angle; opercle 
darker than remainder of head, its inner surface dusky-black. Right side 
mainly cream-white; opercle largely dusky; occasionally with a few widely 
scattered brown melanophores about mouth and near dorsal margin 
of snout. Apparently with little variation in general coloration between 
specimens from differing habitats, but ground coloration is lighter and 
blotches fewer in juveniles (Fig. 5). 

Except where noted, the foregoing description is based on the holo- 
type. 

Distribution: A western Atlantic species presently known from the 
Gulf of Mexico shore and various estuarine habitats on the coast of 
Mexico from Boca del Rio, Veracruz, south to Zacatal, Campeche. Speci- 
mens have been collected from depths of 0-2 meters, over substrates 
ranging from fine sand to dark brown mud. Salinity of 14.8 %c and water 
temperature of 30.1 °C were recorded for the collection of 5 June 1968 
(GCRL2733). 

Etymology: I take pleasure in naming this species after my colleague. 
Dr. Walter Abbott, who assisted in the collection of the type series. 

Reversal: A dextral juvenile (Fig. 5), collected near Boca del Rio 
(GCRL 2733), is of particular interest in that it is the first known in- 
stance of reversal in Citharichthys. This fish ( 15.9 mm SL) has 76 dorsal 
and 54 anal fin rays, 22 caudal vertebrae and exhibits no other abnormal- 
ities. Gudger ( 1935 ) did not include Citharichthys in his review of re- 
versal in flatfishes and there have apparently been no subsequent records. 
Noting a disparity between records of reversal in young and adults of the 
plaice {Pleuronectes platessa), Gudger suggested that mortality of re- 
versed young "approaches 100 percent." A similar situation may pertain in 
Citharichthys or reversal may be extremely rare in both young and adults. 

Discussion and Comparisons: Three species of shallow water Cithar- 
ichthys are presently recognized as occurring in the Gulf of Mexico or 
Caribbean Sea. C. spilopterus (Fig. 6) occurs from New Jersey to 
Brazil, C. arenaceus (Fig. 6) from the West Indies to Brazil and C. uhleri 
( Jordan ) Jordan and Goss which is known only from the unique holotype 
from Haiti. All are similar in general morphology, existing descriptions fail 
to provide adequate means for their separation and identification of speci- 



FiG. 3. Semidiagrammatic delineations of upper jaw dentition (left 
side uppermost). Upper: Citharichthys ahbotti. Middle: 
C. spilopterus. Lower: C. arenaceus. 



364 Proceedings of the Biological Society of Washington 














m'^ ' 



/S 











New Gulf of Mexico flatfish 



365 




f>§'/i'~^'^:^' \, 




Fig. 5. Juvenile Cii/ian'c/ii/iys aZ?&o«i ( GCRL 2733 ) . Upper: Typical, 
sinistra], specimen (15.5 mm SL). Lower: Anomalous, dex- 
tral, specimen ( 15.9 mm SL ) . 

mens from tropical or subtropical waters may be difficult. To these I 
have now added C. abbotti which is similar to both C. spilopterus and C. 
arenaceus and sympatric with the former along the Mexican Gulf Coast. 
I have not examined C. uhleri but the reported combination of 68 dor- 
sal and 52 anal rays, 17-18 gill rakers, 52+ lateral line scales, narrow 
interorbital, dark brown coloration and 21 caudal vertebrae (personal 
count from radiograph) clearly separates this species from both C. abbotti 
and C. spilopterus. Although C. uhleri may eventually prove to be the 
senior synonym of C. arenaceus, it will not be considered further here. 



Fig. 4. Semidiagrammatic delineations of lower jaw dentition (left 
side uppermost). Upper: Citharichthys abbotti. Middle: C. 
spilopterus. Lower: C. arenaceus. 



366 Proceedings of the Biological Society of Washington 





Fig. 6. Upper: Citharichthys spilopterus Giinther GCRL 2776; 115.0 
mm SL. Lower: C. arenaceus Evermann and Marsh USNM 
203510: 120.0 mm SL. 



Fig. 7. Lateral view of left dentary in three species of Citharichthys 
( 110-115 mm SL); indicated teeth are those usually visible in 
this aspect. Upper: ahhotti. Middle: spilopterus. Lower: 
arenaceus. 



New Gulf of Mexico flatfish 



367 






368 Proceedings of the Biological Society of Washington 

Table 5. Observed ranges and means (x) of selected proportional 

measurements of Cithanchthys abbotti (27.6-142.6 mm), C. arenaceus 

(32.6-121.6 mm) and C. spilopterus (39.2-132.0 mm) in percent of 

standard length or head length (HL). 







C. abbotti 






C. arenaceus 




C. spilopterus 






Range 






Range 






Range 


Character 


N 


% SL 
or HL* 


X 


N 


% SL 
or HL* 


X 


N 


% SL 
or HL* X 


Body depth 


35 


44.1-51.4 


47.7 


29 


44.9-57.4 


49.3 


50 


40.4-49.9 45.6 


Caudal peduncle 
depth 


35 


11.7-14.4 


12.9 


29 


12.3-15.8 


13.3 


49 


11.0-13.9 12.8 


Left pectoral 
fin length 


33 


12.3-15.0 


13.8 


24 


12.0-15.4 


13.8 


50 


13.8-17.0 15.5 


Left pelvic 
fin length 


31 


9.7-13.0 


11.7 


28 


7.7-12.8 


10.6 


49 


8.4-11.0 9.9 


Head length 


33 


27.1-31.3 


29.3 


29 


27.5-32.2 


28.9 


49 


26.3-30.9 28.1 


Postorbital length* 


33 


57.6-66.4 


62.3 


29 


59.2-67.7 


64.4 


49 


60.3-71.3 64.7 


Diameter of orbit* 


33 


17.0-23.9 


20.9 


29 


17.3-23.3 


19.6 


47 


13.4-24.8 20.0 


Snout length* 


33 


18.4-21.0 


19.8 


29 


17.4-20.2 


18.7 


43 


17.6-22.1 19.8 


Upper jaw length* 


33 


34.7-39.4 


37.0 


29 


39.1-43.5 


40.7 


48 


35.2-40.1 37.4 


Lower jaw length* 


33 


41.7-48.2 


44.1 


29 


45.7-50.5 


47.9 


44 


41.0-47.2 44.2 


Length of 1st 
dorsal ray* 


29 


23.9-31.0 


27.6 


27 


21.6-26.5 


23.8 


— 


— — 



Fixed teeth of C. abbotti are slightly more numerous than those of C. 
arenaceus and C. spilopterus, as well as somewhat more regular and, pos- 
terolaterally, more strongly recurved (Figs. 3 and 4). Anterior teeth of 
both C. arenaceus and C. spilopterus are prominently enlarged and more 
widely spaced than in C. abbotti; those of C. arenaceus are caniniform 
and set in elevated bony sockets. The left premaxillary teeth are con- 
tinued further posteriad in C. abbotti. Movable teeth are longer in C. 
arenaceus and are proportionately smaller and less abundant in C. spilop- 
terus. The tooth bearing margin of the left dentary is similar in C. ab- 
botti and C. spilopterus but longer and less strongly convex in C. aren- 
aceus (Fig. 7). 

Meristic characters ( Tables 1-4 ) all overlap to some extent but a high 
degree of species separation is indicated. Analyses of variance show ap- 
parent differences to be significant beyond the 1.0 percent level in all 
but three instances. The number of caudal vertebrae is similar in C. 
abbotti and C. arenaceus but different from C. spilopterus. There is no 
significant difference in number of lateral line scales of C. abbotti and C. 
spilopterus although these differ from C. arenaceus. Similarly, there is no 
difference in number of lower arch gill rakers of C. spilopterus and C. ar- 
enaceus but both are significantly different from C. abbotti in this charac- 
ter. Compared with C. arenaceus, C. abbotti averages three more dorsal 
rays, two more anal rays, three less lateral line scales and about three more 



'New Gulf of Mexico flatfish 369 

gill rakers. Compared with C. spilopterus, it averages two less dorsal and 
three less anal rays, four more gill rakers and one less caudal vertebra. 

Proportional measurements also overlap to a considerable degree but in 
most cases the small observed differences show a high order of signifi- 
cance. The range and means of 11 proportional characters are shown in 
percent of SL or HL (Table 5). Serial measurements were not made of 
the 1st dorsal ray in C. spilopterus but measurements of a few specimens 
show it to be short, similar in length to that of C. arenaceus. Results of 
covariance analyses between C. abbotti and the other two species, to- 
gether with regression coefficients and other pertinent data are shown in 
Table 6. 

Right side narial diameters are somewhat greater in C. spilopterus and 
the 1st dorsal ray is inserted above the posterior naris (Fig. 2). Narial 
diameters of C. arenaceus approximate those of C. abbotti and dorsal fin 
origin appears to be intermediate between that of C. spilopterus and its 
anterior location in C. abbotti. 

Citharichthys abbotti is distinct from its compared congeners in a num- 
ber of meristic and morphometric characters. In some features, such as 
numbers of vertical fin rays, it is intermediate between C. arenaceus and 
C. spilopterus. It is inseparable from C. arenaceus on the basis of such 
characters as vertebral number and pectoral fin length and from C. 
spilopterus in such features as jaw lengths, number of lateral line scales, 
etc. C. abbotti is as distinct from each of these species as they are from 
each other and I consider it to warrant specific status. Characters most 
useful in the identification of these forms have been compared in Table 7. 

Although presently known only from the southwestern Gulf of Mexico, 
C. abbotti is likely to occur in inshore and estuarine habitats along the 
Caribbean coast of Mexico and northern Central America. 

It should be noted that current literature recognizes only a single row 
of teeth in the upper and lower jaws of Citharichthys. The keys of 
Norman (1934) and Gutherz (1967) separate Citharichthys from 
Syacium Ranzani on the basis that upper jaw teeth are uniserial in the 
former and biserial in the latter. Whereas this is true for the fixed teeth, 
at least Syacium gunteri Ginsburg and Cyclopsetta chittendeni Bean, a 
related form with uniserial fixed teeth, also have two or three rows of 
movable teeth in each jaw. Since movable teeth can be seen by careful 
examination under low magnification, a definite possibility exists for 
misidentification of these genera. Future keys and descriptions should 
clearly distinguish between fixed and movable teeth. 

Other Material: In addition to listed specimens of C. abbotti, the fol- 
lowing material was examined in this study: Citharichthys arenaceus: 
USNM 49536; holotype; 132 mm SL; Mayaguez, Puerto Rico. USNM 
50195; paratypes; (2) 99-100 mm SL; Mayaguez, P. R. USNM 126424; 
paratypes; (9) 47-103 mm SL; Mayaguez, P. R. USNM 203510; (31) 
27-122 mm SL; Dominica, BWI, mouth of Blenheim River; 7 Novem- 
ber 1964; V. G. Springer and R. H. Reckenveg coll. GCRL 2832; 95 mm 



370 Proceedings of the Biological Society of Washington 

Table 6, Means (x, y), standard deviations (ax, ay), correlation coef- 
ficients (r), standard error (Sy) and regression coefficients (by.x) for 
11 measurements against SL or HL (*) in Citharichthys abbotti, C. 
arenaceus and C, spilopterus. F values measure differences between 
C abbotti and congeners after adjustment for differences between stan- 
dard lengths of samples. Regression equations are in the form: 
y = y + b(x-x). 



Character 
F value 



Species 



N 



Sy 



by.x 



Body depth 
F = 11.8102 
F = 28.4522 

Caudal peduncle 

depth 

F = 6.8051 

F = 2.708 

Left pectoral 
length 
F= 0.008 
F = 73.9872 

Left pelvic length 
F = 19.0292 
F = 81.1172 

Head length 
F= 1.030 
F = 10.1922 

Postorbital 
length* 
F = 31.9842 
F= 9.8212 

Diameter of 
orbit* 
F = 20.930 
F= 0.005 

Snout length* 
F = 47.4102 
F= 0.662 

Upper jaw 
length* 
F = 134.3732 
F = 0.052 

Lower jaw 
length* 
F = 137.4412 
F = 0.809 

1st dorsal ray 
length 
F = 85.3942 

1P< .05; 



abbotti 35 71.8 33.6 3.4 16.5 0.9968 1.3201 0.4897 

arenaceus 29 70.7 20.1 3.5 11.5 0.9932 1.3680 0.5688 
spilopterus 50 87.8 26.7 4.0 13.2 0.9931 1.5624 0.4918 



abbotti 35 71.8 33.6 9.4 

arenaceus 29 70.7 20.1 9.5 
spilopterus 49 87.4 26.9 11.4 



abbotti 

arenaceus 

spilopterus 

abbotti 

arenaceus 

spilopterus 



33 72.3 34.4 10.0 
25 73.4 19.8 10.1 
50 87.8 26.7 13.6 



31 72.9 34.8 
28 71.1 20.3 
50 88.7 26.2 



8.4 

7.5 



4.7 0.9962 0.4110 0.1384 

3.0 0.9886 0.4657 0.1495 

3.8 0.9912 0.5009 0.1384 

4.8 0.9946 0.5072 0.1391 

3.1 0.9826 0.5800 0.1520 
4.0 0.9815 0.7805 0.1482 

4.0 0.9839 0.7186 0.1115 

2.2 0.9695 0.5502 0.1049 
2.6 0.9797 0.5332 0.0984 



abbotti 33 71.1 32.8 20.6 9.0 0.9982 0.5463 0.2754 

arenaceus 29 70.7 20.1 20.3 5.5 0.9900 0.7930 0.2722 
spilopterus 49 88.7 26.1 24.9 7.2 0.9915 0.9529 0.2757 



abbotti 

arenaceus 

spilopterus 

abbotti 

arenaceus 

spilopterus 

abbotti 

arenaceus 

spilopterus 

abbotti 

arenaceus 

spilopterus 

abbotti 

arenaceus 

spilopterus 

abbotti 
arenaceus 

2P< .01. 



33 20.6 9.0 13.0 6.0 0.9991 0.2654 0.6681 

29 20.3 5.5 13.2 3.8 0.9984 0.2231 0.6961 

49 24.9 7.2 16.2 5.2 0.9961 0.4628 0.7174 

33 20.6 9.0 4.2 1.5 0.9921 0.1895 0.1627 

29 20.3 5.5 3.9 0.9 0.9871 0.1497 0.1641 

47 24.7 7.2 4.8 1.1 0.9331 0.3984 0.1411 

33 20.6 9.0 4.1 1.9 0.9980 0.1183 0.2051 

29 20.3 5.5 3.8 1.1 0.9914 0.1461 0.1973 

43 25.6 6.5 5.0 1.2 0.9703 0.2826 0.1722 



33 20.6 9.0 7.7 3.5 0.9974 0.2564 0.3819 
29 20.3 5.5 8.3 2.3 0.9970 0.1843 0.4151 
48 24.8 7.3 9.3 2.8 0.9925 0.3406 0.3728 



33 20.6 9.0 9.0 4.0 0.9977 0.2738 0.4403 

29 20.3 5.5 9.7 2.6 0.9964 0.2267 0.4750 

44 25.5 6.4 11.3 2.9 0.9915 0.3820 0.4465 

33 73.2 34.0 5.9 2.6 0.9907 0.3560 0.0750 

27 70.6 20.3 4.8 1.3 0.9766 0.2911 0.0639 



Islew Gulf of Mexico flatfish 



371 



Table 7. Comparison of external characters most useful in distinguish- 
ing Citharichthys abbotti, C. arenaceus and C. spilopterus. Means and 
standard deviations of meristic characters are in parentheses. 



Character 



abbotti 



arenaceus 



spilopterus 



Dorsal fin rays 
Anal fin rays 
Upper arch 

gill rakers 
Total gill rakers 
Number of lateral 

line scales 
Anterodorsal 

contour 
Left pectoral 

fin length 
Left pelvic 

fin length 
Lower jaw length 
Length of 1st 

dorsal ray 
Color in alcohol 



72-81 (76 ± 1.7) 

52-60 (56 ± 1.5) 

5-8 (6 ± 0.7) 

18-23 (21 ±1.1) 
42-49 (45 ± 1.5) 

Convex, slight notch 

before eye 
averages 14% of SL 



70-77 (73 ± 1.6) 

51-56 (54 ± 1.3) 

3-7 (5 ± 0.8) 

15-21 (18 ±1.2) 
45-50 (48 ± 1.3) 

Convex, distinct 

notch before eye 
similar to abbotti 



averages 12% of SL averages 11% of SL 



averages 44% of HL 
averages 28% of HL 

Light sandy-tan, 
sprinkled with 
brown flecks 



averages 48% of HL 
averages 24% of HL 

Dark brown with 
tan or light 
brown flecks 



74-82 (78 ± 1.8) 

56-62 (59 ± 1.5) 

3-5 (4 ± 0.6) 

13-19 (17 ±1.3) 
43-48 (45 ± 1.3) 

usually concave 

averages 15.5% of SL 

averages 10% of SL 

similar to abbotti 
similar to arenaceus 

sandy-brown to gray, 
usually with some 
brownish flecks 
on fins 



SL. GCRL 2810; (2) 107-111 mm SL (cleared and stained). Data 
for GCRL specimens as for USNM 203510. 

Citharichthys spilopterus: Northern Gulf of Mexico — GCRL 2776; 
(15) 82-117 mm SL; 29°42'N, 88°27'30"W; 60 meters; 17 January 1968; 
R/V GULF RESEARCHER coll. GCRL 2795; (17) 93-113 mm SL; lo- 
cation as for preceding collection; April- June 1967; R/V GULF RE- 
SEARCHER coll. GCRL uncataloged; (21) 89-127 mm SL (two cleared 
and stained ) ; other data as for GCRL 2795. 

Mexico, Veracruz — GCRL 2367; 128 mm SL; south jetty at Alvarado; 
3 September 1967; C. E. Dawson coll. GCRL 2686; (2) 46-62 mm SL; 
north shore of Laguna Zontecomapan, about 1/4 mi. from mouth; 1 Sep- 
tember 1967; C. E. Dawson and W. Abbott coll. GCRL 2687; (5) 39-132 
mm SL; south side of Laguna Zontecomapan, about 3/4 mi. from mouth; 
1 September 1967; C. E. Dawson and W. Abbott coll. GCRL 2726 (7) 
21-85 mm SL; Rio Jamapa, southwest end of bridge on Mexico 180; 5 
June 1968; C. E. Dawson and S. Romay coll. GCRL 2734; (16) 40-103 
mm SL; Isla da Amor, Boca del Rio; 5 June 1968; C. E. Dawson coll. 
GCRL 2833; (2) 59-73 mm SL; tidepool on north side of south jetty 
at Alvarado; 7 June 1968; C. E. Dawson coll. GCRL 2806; (2) 30-33 
mm SL; north shore of Isla Pajaritos, Rio Coatzacoalcos; 8 June 1968; C. 
E. Dawson coll. GCRL 2745; 48 mm SL; mouth of Laguna Zontecoma- 
pan; 16 August 1968; C. E. Dawson coll. 



372 Proceedings of the Biological Society of Washington 

Acknowledgments: Appreciation is expressed to Dr. V. G. Springer 
and Robert Kanazawa ( USNM ) for the loan and exchange o£ specimens 
of C. arenaceus and for radiographs of pertinent type material in the 
National collections. Mrs. M. M. Dick (MCZ) kindly provided a radio- 
graph of the holotype of C. uhleri and Sr. Humberto Chavez, Direccion 
General de Pesca e Industrias Conexas, has been most helpful in expedit- 
ing my fieldwork in Mexico. Mr. A. C. Wheeler, British Museum ( Nat- 
ural History), is thanked for providing data on the holotype of Cithar- 
ichthys spilopterus. Drawings are by Mr. Harry L. Moore, Jr. and Mr. 
Charles Eleuterius provided computer programs and statistical analyses. 

This study was in part supported by National Science Foundation 
Grant GB-6823. 

Literature Cited 

GuDGER, E. W. 1935. Abnormalities in flatfishes ( Heterosomata ) . I. 
Reversal of sides — A comparative study of known data. J. 
Morph. 58(1): 1-39. 

GuTHERz, E. J. 1967. Field guide to the flatfishes of the family Bothidae 
in the western north Atlantic. U. S. Fish Wildl. Serv. Cir- 
cular 263: 1-47. 

Norman, J. R. 1934. A systematic monograph of the flatfishes ( Hetero- 
somata), Vol. I, Psettodidae, Bothidae, Pleuronectidae. Brit. 
Mus. (Nat. Hist.), vii + 459 pp. 

Parr, A. E. 1931. A practical revision of the western Atlantic species of 
the genus Citharichthys (including Etropus). Bull. Bingh. 
Oceanogr. Coll. 4(1): 1-24. 



GH 

i 

B4X 



7^ 



SO 3 October 1969 



PROCEEDINGS 
OF THE 
BIOLOGICAL SOCIETY OF WASHINGToti OCT 20 1969 'i 



PARAGUNNELLICHTHYS FEHLMANNI, A NEW 

GOBIOID FISH (MICRODESMIDAE) FROM THE 

INDIAN OCEAN 

By C. E. Dawson 

Gulf Coast Research Laboratory, 

Ocean Springs, Mississippi 

Among recently received collections of Indo-Pacific worm- 
fishes there is an undescribed species exhibiting the first known 
occurrence of fully united pelvic fins within the family Micro- 
desmidae. In continuance of my studies of microdesmids, I 
herewith describe this unusual form from the central Indian 
Ocean. 

Specimens reported here were made available for study by 
the Smithsonian Oceanographic Sorting Center and have been 
deposited in the collections of the Smithsonian Institution 
(USNM) and the Gulf Coast Research Laboratory (GCRL). 
Head length is measured from the tip of the lower jaw to base 
of the uppermost pectoral ray; body depth is measured at anal 
fin origin. Caudal fin length is the distance from the rear of the 
hypural to the tip of the longest ray. All fin-rays are counted 
separately; vertebral counts are from radiographs or cleared 
and stained specimens. 

Appreciation is expressed to Dr. Daniel M. Cohen for com- 
ments on the manuscript. Semidiagrammatic delineations are 
by Mr. Harry L. Moore, Jr. This study was supported in part by 
National Science Foundation Grant GB-6823. 

Paragunnellichthys fehlmanni new species 

Holotype: USNM 203830; 33.6 mm SL; Diego Garcia Atoll, Chagas 
Archipelago, 7°15'43"S, 72°22'36"E; chemical ichthyocide collection in 
depths up to 8 ft over reef NW of lagoon; 18 June 1967; H. A. Fehlmann 
coll. 

27— Proc. Biol. Soc. Wash., Vol. 82, 1969 (373) 



CT201969 



374 Proceedings of the Biological Society of Washington 






Fig. 1. Paragunnellichthys fehlmanni. USNM 203830; holotype; 33.6 
mm SL. 



Paratypes: USNM 203832; (2) 31.5-33.8 mm SL. USNM 203831; 
24.9 mm SL; cleared and stained. GCRL 3452; 31.2 mm SL; cleared and 
stained. Collection data of paratypes as for holotype. 

Diagnosis: A species of Paragunnellichthys with short pelvic fins united 
by a membrane; interradial membranes of dorsal and anal fins with a some- 
what irregular series of vertically elongate black blotches; the first two 
dorsal spines more closely spaced than successors; anal fin origin beneath 
interspace between 22nd to 24th dorsal fin elements; proximal pterygio- 
phore of 1st dorsal spine inserted between 3rd and 4th abdominal verte- 
brae. 

Description: Dorsal spines 19-21, dorsal segmented rays 31-33, total 
dorsal elements 51-53; anal rays 29-31; pectoral rays 10; pelvic fin I, 2; 
segmented caudal rays 15; vertebrae 25 + 27-29 = 52-54. See Table 
1 for proportional measurements and counts. 



\ 



New Indian Ocean gobioid fish 



375 



Table 1. Measurements (mm) and counts of Paragunnellichthys 
fehlmanni. 



] 


USNM 
203830 
Holotype 


USNM 
203832 
Paratype 


USNM 
203832 
Paratype 


USNM 
203831 
Paratype 


GCRL 

3452 
Paratype 


Standard length 


33.6 


33.8 


31.5 


24.9 


31.2 


Caudal fin length 


3.6 


3.5 


3.4 


2.6 


3.0 


Least caudal peduncle depth 


1.4 


1.2 


1.2 


1.0 


1.2 


Body depth at anal fin origin 


2.1 


2.0 


2.0 


1.7 


1.9 


Predorsal length to 
tip of lower jaw 


5.1 


5.0 


4.9 


4.2 


4.9 


Preanal length 


18.2 


17.8 


16.8 


13.4 


17.5 


Pectoral fin length 


2.2 


2.1 


2.1 


1.9 


2.2 


Pelvic fin length 


1.1 


0.9 


1.0 


0.8 


0.9 


Distance from pelvic insertion 
to anal fin origin 


13.7 


13.4 


12.8 


9.5 


13.3 


Head length 


4.2 


4.1 


3.7 


3.4 


4.1 


Eye diameter 


0.6 


0.6 


0.6 


0.5 


0.6 


Snout length 


0.6 


0.6 


0.5 


0.5 


0.6 


Distance from anterior margin 
of eye to tip of lower jaw 


0.9 


0.9 


0.9 


0.8 


1.0 


Postorbital length 


2.6 


2.6 


2.2 


2.1 


2.5 


Interorbital width 


0.3 


0.3 


0.3 


0.2 


0.3 


Tip of lower jaw 
to angle of gape 


0.9 


0.7 


0.8 


0.7 


0.7 


Number of dorsal spines 


20 


20 


21 


19 


20 


Number of segmented 
dorsal rays 


31 


32 


32 


33 


32 


Total dorsal elements 


51 


52 


53 


52 


52 


Number of anal rays 


29 


30 


31 


30 


29 


Anal fin origin beneath 
interspace between 
dorsal elements 23/24 22/23 23/24 22/23 23/24 



Body moderately elongate, slender, depth at anal fin origin averages 
6.3 percent of SL, tapering to about 60 percent of body depth at caudal 
peduncle; compressed, breadth at anal fin origin 2.7 percent of SL in 
holotype, greatest breadth (5.1 percent of SL) at opercle; caudal fin 






376 Proceedings of the Biological Society of Washington 




Fig. 2. Paragunnellichthys fehlmanni. 
tion of dorsal aspect of head. 



Semi-diagrammatic delinea- 



broadly rounded, averages 10.4 percent of SL; head length 11.7-13.7 per- 
cent of SL, its depth subequal to that of body; interorbital narrow, convex, 
its width equal to about 1/2 of eye diameter; eye lateral, high on head, 
its diameter, averaging 14.9 percent of head length, is slightly greater 
than snout length; lower jaw prominent, extends about 2/3 of eye diam- 
eter beyond snout tip, fleshy, its lateral depth subequal to eye diameter, 
narrowing in front to a prominent subvertical fleshy symphysial ridge 
which is matched by a slight emargination of the snout tip ( Fig. 2 ) ; gape 
short, reaches a vertical from front margin of posterior naris, inclined to 
about 45°; upper lip broad near angle of gape, narrow and concealed 
in front by overhanging snout; lower lip broad posteriad but 
narrows to a faint ridge across the symphysis, somewhat pouch-Hke 
and includes the anterior 2/3 of the upper lip and the median snout tip 
when mouth is closed; anterior naris opens anterolaterally through a 
short tubule at the end of a prominent fleshy dorsolateral longitudinal 
snout ridge; posterior naris dorsolateral over preorbital, with only a 
slightly elevated margin, its diameter about 7 in eye; lower jaw with 
two anterior rows of 12-14 large caniniform teeth and with a single row 
of minute pointed teeth on the posterior half of each dentary; upper jaw 
with two rows of about 18-20 enlarged caniniform teeth, those of the 
inner row somewhat recurved. 



( 



New Indian Ocean gobioid fish 377 




Fig. 3. ParagunneUichthys fehlmanni. Semi-diagrammatic delineation 
of united pelvic fins. 

Gill opening large, subtubiform, directed posteriad, originates on pec- 
toral peduncle just anteriad of insertion of uppermost pectoral ray, 
curves dov^nward and slightly forward to unite with the ventral margin 
of the peduncle near its insertion; ventral terminus of gill opening con- 
cealed by folds of the overhanging branchiostegal membrane. Dorsal and 
anal rays terminate on the short caudal peduncle, free from the caudal 
fin, depressed tips of terminal rays extend slightly past rear of hypural; 
dorsal fin originates well posteriad of gill opening, near a vertical through 
anterior third of pectoral fin; interspace between the 1st two dorsal 
spines one-third to one-half shorter than those which follow; the 1st 
dorsal spine about two-thirds the length of the 2nd, the 1st segmented ray 
is about 20 percent longer than the last spine; the last two dorsal and 
anal rays separate, not closely approximated; the 1st anal element is 
segmented and all segmented dorsal and anal rays are apparently simple; 
caudal fin formula, from stained paratype, 5 + 2 + 11 + 2 + 5; pectoral 
fin elongate and narrowly rounded, fin-rays simple, the 5th or 6th the 
longest, the rather short peduncle is not completely concealed by the 
opercle and branchiostegal membrane. Pelvic fins (Fig. 3) inserted 
about one-half their length in advance of dorsal fin origin and 
well posteriad of pectoral fin insertion, united by a distinct membrane, 
each with a slender outer spine and two simple rays, the innermost the 
longer. Anal fin origin beneath interspace between dorsal elements 22 
and 23 or 23 and 24. 

Head with embedded, separated, cycloid scales on posterior nape, 
opercle and upper branchiostegal membrane, apparently absent from 
cheeks, snout and ventral head; body scales similar, somewhat larger, 
nonimbricate but occasionally touching, cover predorsum, lateral body 
and abdomen except for a small naked area about anal and genital open- 
ings; fins naked but with some scales on pectoral peduncle and con- 
tinuing forward within the gill opening; scales inconspicuous and dif- 
ficult to see, maximum scale diameter about four in eye diameter. 



378 Proceedings of the Biological Society of Washington 



Fig. 4. Paragunnellichthys fehlmanni. Semi-diagrammatic delineation 
of median body section showing dorsal and anal fin blotches. 

Dermal sensory papillae inconspicuous, apparently with a transverse 
row crossing nape, a short longitudinal row along upper margin of oper- 
cle, a supraorbital row which continues forward to snout tip, one or 
two rows on infraorbital and lower preorbital and with two parallel 
rows along side of lower jaw; sensory papillae apparently absent from 
abdomen and lateral body. 

Apparently without predorsal interneurals; pterygiophores poorly os- 
sified and difficult to see in both radiographs and stained material; 
distal pterygiophores begin immediately behind the first segmented dor- 
sal ray and after the second anal ray; first dorsal spine inserted over inter- 
space between the 3rd and 4th abdominal vertebrae. Abdominal neural 
spines strong, slender and pointed, generally of subequal length and 
located posteriad on centra; prezygopophyses low, slightly elevated and 
bluntly pointed in lateral aspect; ribs apparently articulate with centra 
of the first two vertebrae, with parapophyses on remaining abdominal 
vertebrae; abdominal epipleurals begin with the third vertebra; skull 
short, somewhat elevated, frontals poorly ossified and fail to reach mes- 
ethmoid in either radiographs or stained material; posttemporals curve 
well forward, not elevated above the anteriormost vertebrae; branchios- 
tegal rays 5, the innermost remote from its fellows; mandible short and 
well ossified, with a prominent, pointed, terminal ventral process. 

Ground color, in alcohol, primrose yellow; without conspicuous mark- 
ings on head or body but with a few irregularly scattered light brown 
micromelanophores on predorsum, along either side of the spinous dorsal 
and on the midlateral body; interradial membranes of dorsal and anal 
fins with prominent but somewhat irregular series of vertically elongate 
dark brown to black blotches (Figs. 1 and 4); dorsal markings are more 
closely spaced than those of the anal fin and counts from three fish 
ranged from 23 to 27 on the dorsal membranes and 10 to 13 on the 
anal; remaining interradial membranes, caudal and paired fins immacu- 
late; eye black, the pupil somewhat iridescent. 



I 



New Indian Ocean gobioid fish 



379 



Table 2. Comparison of selected characters distinguishing Paragunnel- 
lichthys fehlmanni and P. seychellensis. 



Character 


P. fehlmanni 


P. seychellensis 


Number of dorsal spines 


19-21 


16^18 


Total dorsal elements 


51-53 


47-48 


Pelvic fins 


united, inserted pos- 
teriad of pectorals 


separate, inserted in 
advance of pectorals 


Number of abdominal 
vertebrae 


25 


21 


Total vertebrae 


52-54 


48-49 


1st dorsal spine inserted 
between 


3rd and 4th 
vertebrae 


1st and 2nd 
vertebrae 


Postorbital length 


averages 61.5 
percent of HL 


averages 53.4 
percent of HL 


Tip of lower jaw to 
angle of gape 


averages 19.6 
percent of HL 


averages 25.4 
percent of HL 



Etymology: I take pleasure in naming this species after the collector, 
Dr. H. A. Fehlmann. 

Relationships: The presence of conspicuous blotches on the dorsal and 
anal fins permits ready separation of Paragunnellichthys fehlmanni from 
its only known congener P. seychellensis Dawson. These species are 
further differentiated in a number of meristic and morphological features 
and some of the more useful characters are compared in Table 2. 

The completely united pelvic fins of P. fehlmanni are unique among 
microdesmids. I have seen a narrow fragile basal membrane uniting these 
fins in one or two of some 13 specimens of Gunnellichthys irideus J. L. 
B. Smith, but separate pelvic fins has formerly been considered to be a 
family characteristic. In the absence of other major anatomical varia- 
tions, I do not consider that the united fin condition justifies erection of 
a new genus. 

In addition to the united pelvics, Paragunnellichthys fehlmanni dif- 
fers from the generic description (Dawson, 1967) in having a more pos- 
terior insertion of both dorsal and pelvic fins. It does agree, however, in 
such critical features as the nature of the gill opening and the number of 
pelvic and principal caudal fin-rays. Despite certain discrepancies, it is 
evident that fehlmanni should be included in Paragunnellichthys in pre- 
ference to the other microdesmid genera which consistently have either 
three or four segmented pelvic rays. 



380 Proceedings of the Biological Society of Washington 

Remarks: Paragunnellichthys is now known only from the Seychelles and 
the Chagas Archipelago. Although this genus may have restricted distribu- 
tion, it should be expected in other Indo-Pacific reef and lagoon habitats. 
This conclusion is supported by the recent collection of Gunnellichthys 
curiosus Dawson, previously known only from the Seychelles, in Hawaiian 
waters by Dr. J. E. Randall. 

Literature Cited 

Dawson, C. E. 1967. Paragunnellichthys seychellensis, a new genus and 
species of gobioid fish (Microdesmidae) from the western 
Indian Ocean. Proc. Biol. Soc. Wash. 80: 73-82. 



1 

B4X 
NH 



3 October 1969 



PROCEEDINGS ^^HS07v^^ 

OF THE /^^ ^4^ 

BIOLOGICAL SOCIETY OF WASHINGT^IsPCT 2 Q 1969 

NEW SPECIES OF SPIONIDS (POLYCHAETA) FROM 

THE GULF OF MEXICO AND CARIBBEAN SEA WITH A 

PARTIAL REVISION OF THE GENUS PRIONOSPIO 

By Nancy M. Foster 
Smithsonian Institution, Washingtony D.C. 

In connection with a study of the Spionidae of the Gulf of 
Mexico and Caribbean Sea, several new taxa have been en- 
countered. This paper deals with the descriptions of two new 
species, one of which represents a new genus, and the restab- 
lishment of a second genus. 

I wish to acknowledge Dr. Olga Hartman and Dr. Lucien 
Laubier for the loan of type material for examination. I thank 
Dr. John Day for the offer of his specimens, notes and draw- 
ings of one of the new species described herein. Finally, I 
would like to thank Dr. Marion Pettibone and Dr. Meredith 
Jones for their constructive comments with regard to the writ- 
ing of this manuscript. This work was supported by a Smith- 
sonian Pre-doctoral Internship and all type material is de- 
posited in the Smithsonian Institution. 

The following are covered in this study; 

Discussion of Prionospio Malmgren, 1867 
Apoprionospio new genus A. dayi new species 

A. nova (Annenkova, 1938) new 

combination 

A. saldanha (Day, 1961) new 

combination 

A. pygmaea (Hartman, 1961) 

new combination 

A. caspersi (Laubier, 1962) new 

combination 

Key to species of Apoprionospio 

2&— Proc. Biol. Soc. Wash., Vol. 82, 1969 (381) 



382 Proceedings of the Biological Society of Washington 

Paraprionospio CauUery, 1914 [= Prio7iospio ( Paraprion- 

ospio) Caullery] 

P. pinnata (Ehlers, 1901) new 

combination 
Aonides Claparede, 1864 A. mayaguezensis new species 

Key to species of Aonides 

The genus Prionospio Malmgren, 1867 was originally limited 
to species possessing gills beginning on setiger two. Caullery 
( 1914 ) erected Paraprionospio as a subgenus to accommodate 
the species Prionospio pinnata, which possesses branchiae 
beginning on the first setiger. Later Paraprionospio was used 
at the generic level by Berkeley ( 1927 ) and Weese ( 1933 ) . This 
usage, however, was not continued by later systematists. Rather 
than accept Paraprionospio as a genus or subgenus, the limits 
of Prionospio were merely expanded to include it. This ap- 
parently has been the tendency with regard to Prionospio with 
the result that at present, Prionospio includes an extremely 
heterogenous group' of species. The species of this genus can be 
separated into distinct groups on the basis of type and arrange- 
ment of the gills. Previously all species with the following 
characters have been assigned to the genus Prionospio: 

1. anterior branchiae first present on setigers one, two or 
three; separate from dorsal lamellae; pinnate and/ or 
cirriform. 

2. rounded prostomium, lacking frontal horns and occipital 
tentacles. 

3. neuropodial and notopodial hooded hooks, bidentate or 
multidentate. 

4. well-developed anterior parapodial lamellae. 

On the basis of the above, the generic characters of Prionos- 
pio do not consider the question of gill type and arrangement, 
giving no indication of their pinnate or cirriform nature. In 
other valid genera of the Spionidae, the species form homogene- 
ous groups with regard to the branchiae. Therefore, after con- 
sideration of the characters involved in generic diagnoses within 
the family as a whole, I have decided to divide Prionospio into 
several genera. It is recognized that these divisions may be 



Two new spionids 383 

found to be too artificial and that subgeneric designations 
would be more acceptable. 

Two groups of species are herein removed from Prionospio 
and placed in Paraprionospio CauUery and Apoprionospio new 
genus. 

Apoprionospio new genus 

Type species: Apoprionospio dayi new species. 

Diagnosis: Prostomium triangular, flared anteriorly, lacking frontal 
horns, bearing two pairs of eyes. Peristomium fused with setiger one 
surrounding prostomium posteriorly as a yoke or collar; with no lateral 
wings or elevations. Four pairs of branchiae beginning on setiger two; 
first three pairs cirriform, fourth pinnate. Neuropodial lamellae of setiger 
two somewhat enlarged, triangular, directed ventrally. Pygidium with 
anal cirri. Hooded hooks in posterior neuropodia and notopodia; biden- 
tate or multidentate. Dorsal membranous ridges connecting notopodial 
lamellae at some point along the body. 

Etymology: Gender — feminine; apo — Gr. — from, away from, sepa- 
rate; priono — <Gr. — saw; spio — L. — a sea nymph; alluding to the separa- 
tion of the members of this genus from Prionospio. 

Apoprionospio dayi new species 
(Figs. 1-11) 

Diagnosis: Body up to 35 mm in length. Triangular prostomium with 
two pairs of eyes and occasionally additional eye spots. Peristomium in 
form of a yoke or collar rather than hood. Four pairs of branchiae; first 
to third, cirriform and fourth, pinnate, with pinnae extending only 1/2 
to 2/3 length of the shaft. Neuropodial lamellae of setiger two larger 
than those of remaining setigers. Dorsal crest, between notopodial lamel- 
lae, on setiger seven. Multidentate, hooded hooks first appearing in seti- 
gers 16 to 18 of neuropodia and in setigers 27 to 40 of the notopodia. 
Pygidium with three anal cirri, one long mid-dorsal and two shorter later- 
als. 

Material examined: off Beaufort, North Carolina (five specimens). 
Grand Isle, Louisiana (one specimen) and Port Aransas, Texas (two 
specimens ) . 

Type Material: Holotype: USNM 39487; Paratype: USNM 39488. 
Collected off Beaufort, North Carolina. 

Etymology: The species is named for Dr. John Day, Capetown, South 
Africa. 

Description: The prostomium (Fig. 1) is triangular in shape, widest 
anteriorly. It narrows posteriorly and extends to about the level of setiger 
two. There are two pairs of eyes. Those of the anterior pair are farther 
apart and may be larger. There is a ridge behind the prostomium which 
extends across the dorsum from just behind the first parapodia. The per- 






384 Proceedings of the Biological Society of Washington 




Two new spionids 385 

istomium is low and surrounds the prostomium like a collar rather than 
the more typical hood. There are no lateral wings. In some specimens 
the transverse ridge appears to be a part of this peristomial collar, its 
posterior continuation. 

There are four pairs of branchiae; the first three cirriform and the 
fourth, pinnate. The first pair is slender and neither obviously ciliated 
nor tapered ( Figs. 1 and 2 ) . The second and third pairs are thicker and 
tapered, with long cilia on the proximal one-half to two-thirds of the gill 
shafts and shorter cilia continuing to the branchial tips (Figs. 1 and 3). 
The pinnate branchiae of setiger five are the longest of the four pairs. 
They are pinnate, the pinnae extending one-half to two-thirds the length 
of the shafts. They are as long as the gill width and closely adhere to one 
another forming one row on either side of the gill ( Fig. 4 ) . 

The anterior notopodia are long and extend well above the body. The 
notopodial lamellae of setiger one are low, rounded and slightly cupped, 
while those of the neuropodia are taller and thinner (Figs. 1 and 5). The 
parapodial lamellae of setiger two are conspicuously different from those 
of the preceeding in that the neuropodial lamellae are especially large 
and triangular in shape with the long axis extending ventrally (Figs. 1 
and 2). Notopodial lamellae have become elongate, erect and triangular 
with a slight anterior fold or cup opening laterally, from which the noto- 
setae emerge. The fold becomes more pronounced in the following 
setigers and by setigers four to five it is almost symmetrical, forming a 
V-shaped cup or groove with both sides equal. At this point the lamellae 
are very elongate (Fig. 3) and neuropodial lamellae are thinner and 
triangularly tapered. 

On setiger seven a dorsal ridge or crest extends across the dorsum 
between the notopodial lamellae. The anterior face of the cup on this 
setiger is more like a presetal lamella and is rounded (Fig. 6). The ridge 
extends from the point of "fusion" of the pre- and postsetal lamellae. The 
neuropodial lamella is bluntly rounded and similar to that of setiger one. 
The neuropodial presetal lamella is fairly well-developed, low and 
rounded. 

On the next few setigers the notopodial lamellae are flattened, low 
and may extend upon the dorsal surface but do not connect to form 
ridges; both noto- and neuropodial presetal lamellae are well-developed 
(Fig. 7). In posterior setigers the notopodial lamellae are much more 
lateral, forming low ridges approximately level with the dorsum; the pre- 
setal lamellae have decreased in prominence and are present as low 
ridges; the setal bundle has shifted to a more ventral position. (Fig. 8). 



Figs. 1-5. Apoprionospio dayi new genus, new species. 1. Dorsal view, 
anterior end. 2. Setiger two, showing enlarged neuropodial lamella. 3. 
Setiger three. 4. Setiger five. Figs. 1-4, scale B; Fig. 5, scale C. 



386 Proceedings of the Biological Society of Washington 




Figs. 6-11. Apoprionospio dayi new genus, new species. 6. Setiger 
seven. 7. Setiger 11. 8. Setiger 22. 9. Neuropodial seta from setiger one. 
10. Sabre-seta from setiger 12. 11. Neuropodial hooded hook from posterior 
setiger. Figs. 12^13. Paraprionospio pirmata. 12. Dorsal view, anterior 
end, bases of gills shown. 13. Setiger one, pigmentation of gills shown by 
stipling. Figs. 9-11, scale A; Figs. 6-8, 13, scale B; Fig. 12, scale D. 



Two new spionids 387 

Neuropodial lamellae are similar in shape to those of the notopodia but 
remain smaller. Posteriorly the notopodial lamellae become taller and 
thinner and those of the neuropodia lower and more rounded. 

Anterior capillary setae are all similar, appearing unilimbate. Current 
work soon to be completed has indicated, however, that what was pre- 
viously considered to be a limbation Is in actual fact a "sheath," partially 
or completely surrounding the setal shaft. In A. dayi the capillaries show 
granulations, or what have previously been termed spinelets (Jones, 
1962), along the area inclosed within the sheath. Two areas are clearly 
distinguishable on the shaft, a smooth clear outer part and a granular inner 
part (Fig. 9). There are few setae in setiger one. In the second setiger 
the notosetae increase significantly in number, and are arranged in a 
whorl and vary considerably in length. By setigers four to five, neuro- 
podial setae are thinner and not as broadly sheathed as those of the 
notopodia. A large sabre-seta appears in setiger 11 on all specimens ex- 
amined. It differs from the typical spionid sabre-seta in that it is very 
broadly sheathed (Fig. 7 and 10). Furthermore the sabre-setae do 
not always occupy the ventral-most position of the neuropodial fascicle 
but are often curved to emerge at the same level as some of the hooded 
hooks (Fig. 8). 

In several setigers anterior to the first appearance of hooded hooks, 
neuropodial setae have become very long and are no longer granulated 
and broadly sheathed. Neuropodial hooded hooks appear in setigers 16 
to 18. The hooks have long primary hoods and very small, closely ap- 
plied secondary or internal hoods. The presence of these internal hoods 
was first suggested by Jones ( 1962 ) and they were referred to as "inter- 
nal cylinders." The hooded hooks have two to three small teeth above 
the main fang (Fig. 11). The distribution of two and three denticled 
hooks varies and there does not seem to be a pattern. The neuropodial 
fascicle also includes long, thin, companion setae ( Fig. 8). Notopodial 
hooded hooks, similar to those of the neuropodia, first appear in setigers 
27 to 40. 

The pygidium bears a single, long, slender, mid-dorsal and two shorter, 
lateral anal cirri. 

Discussion: According to the above revision, the following species of 
Prionospio may be referred to Apoprionospio new genus: 

A. nova (Annenkova, 1938) new combination. Japan Sea. Indeter- 
minable. 

A. saldanha ( Day, 1961 ) new combination. South Africa. 

A. pygmaea (Hartman, 1961) new combination. Southern Cali- 
fornia. 

A. caspersi (Laubier, 1962) new combination. Lagune de Venice, 
Adriatic. 

A. dayi new species. North Carolina, Gulf of Mexico (Louisiana, 
Texas). 



388 Proceedings of the Biological Society of Washington 

Apoprionospio dayi differs from the closely related A. pygmaea by the 
presence of the dorsal membranous ridge on setiger seven. Examination 
of type material of the latter, deposited in the Allen Hancock Foundation, 
revealed the presence of multidentate hooded hooks rather than bidentate 
as indicated in the original description. The pinnate branchiae of A. dayi 
differ from those of A. caspersi and A. saldanha by having the pinnae 
regularly arranged in two opposing rows along the shaft, closely applied 
to one another and extending only about one-half the length of the gill. 
Examination of paratypes of the above confirmed the irregular arrange- 
ment of the pinnae and their extension along almost the entire length of 
the shaft. A. pygmaea and A. dayi possess multidentate hooks whereas in 
A. caspersi and A. saldanha they are bidentate. Priorwspio nova, based on 
an incomplete specimen, is considered indeterminable at the species level; 
however, the gill arrangement and shape of the second neuropodial lamella 
agree with Apoporionospio. 

KEY TO THE SPECIES OF THE GENUS APOPRIONOSPIO 

1. Neuropodial hooded hooks multidentate. Pinnae of fourth bran- 
chial pair regularly situated in two opposing rows, not extending 
to the end of the shaft 2 

1. Neuropodial hooded hooks bidentate. Pinnae irregularly arranged 

along shaft, extending to near tip of gill shaft 3 

2. Dorsal membranous ridge on setiger seven present A. dayi 

2. Dorsal membranous ridge on setiger seven absent A. pygmaea 

3. Dorsal ridge on setiger seven present A. caspersi 

3. Dorsal ridge on setiger seven absent A. saldanha 

Genus Paraprionospio Caullery, 1914 

Etymology: Gender: — feminine; para — Gr. — beside, near, by. 

Type Species: Prionospio pinnata Ehlers, 1901, designated by Caul- 
lery, 1914. 

Diagnosis: Prostomium surrounded by well-developed peristomial hood 
forming lateral wings. Branchiae pinnate, beginning on setiger one. 
Parapodia of first setiger well-developed, not particularly reduced. Noto- 
and neuropodial hooded hooks multidentate. Pygidium with anal cirri. 

Remarks: Paraprionospio was originally erected for the species com- 
monly known as Prionospio pinnata. In Prionospio sensu stricto the first 
setiger (not segment) is reduced and does not bear the gills; segments 
one and two are fused resulting in the loss of the first parapodia. In P. 
pinnata, however, setiger one is only slightly smaller and does bear the 
first pair of gills. For this reason, Caullery (1914) established the sub- 
genus Paraprionospio. He pointed out that possibly either the first setiger 
was simply not reduced as in Prionospio or that the first parapodia have 
disappeared, with the lateral wings of the head remaining as vestiges of 
the first setiger. Later Soderstrom (1920) synonymized the two genera 



Two new spionids 



389 



saying that, as in Prionospio, segments one and two have fused resulting 
in parapodial loss so that the gills actually begin on segment three. The 
fact remains, however, that if one considers a suite of characters, Par- 
aprionospio can be distinguished as a separate genus. The following Table 
illustrates some of these differences. 

Table 1. Comparison of Paraprionspio and Prionospio. 



First 

appearance 

of gills 



Peristomial 
wings 



Size of 

first 

parapodia 



Paraprionospio setiger 1 



very 
pronounced 



Prionospio 



setiger 2 



not 

significantly 

reduced 

much 
reduced 



Paraprionospio also shows a very close relationship to the genus Stre- 
hlospio. Characteristics in common include prostomium enclosed in a 
hood, branchiae beginning on setiger one and similar setae. 

Paraprionospio pinnata (Ehlers, 1901) new combination 
(Figs. 12-21) 

Prionospio pinnata Ehlers, 1901, p. 163; 1912, p. 110-Augener, 1927, 
p. 351.-Monro, 1923, p. 68.-Fauvel, 1936, p. 60-Okuda, 1936, p. 
247; 1937, p. 49.-Berkeley and Berkeley, 1941, p. 42; 1952, p. 30.- 
Wesenberg-Lund, 1949, p. 325.-Hartman and Reish, 1950, p. 29.- 
Tebble, 1955, p. 124.-Hartman, 1955, p. 182; 1963a, p. 43; 1963b, 
pp. 74, 123, 131, 148, 167, 179, 194, 211, 231, 247, 258, 277, 296, 
328, 344, 358, 375, 398; 1967, pp. 11, I13.-Kirkegaard, 1959, p. 
22.-Reish, 1961, p. 86; 1959, pp. 38, 61, 64, 67, 71, 75, 78.-Bar- 
nard and Reish, 1959, pp. 9, 71, 73, 88.-Day, 1961, p. 485; 1967, p. 
488.-WU and Chen, 1963, p. 59.-Bellan, 1964, p. I12.-Shepherd, 
1964, p. 71.-Imajima and Hartman, 1964, p. 286.-Hartmann- 
Schroder, 1965, p. 211.-Guille and Laubier, 1966, p. 272.-Est- 
court, 1967, p. 76.-Banse and Hobson, 1968, p. 29. 

Prionospio {Paraprionospio) pinnata inaequibranchia CauUery, 1914, 
p. 356. 

Prionospio africana Augener, 1918, p. 402.-Monro, 1930, p. 149. 

Prionospio alata Moore, 1923, p. 185. 

Paraprionospio tribranchiata Berkeley, 1927, p. 415.-Weese, 1933, pp. 20', 
21. 

Prionospio plumosa Treadwell, 1931, p. 4. 

Prionospio tenuis, Hartman, 1945, p. 32. 

Prionospio treadwelli Hartman, 1951, p. 84. 

Prionospio ornata Berkeley and Berkeley, 1961, p. 660. 



390 Proceedings of the Biological Society of Washington 




Figs. 14-21. Paraprionospio pinnata. 14. Setiger eight. 15. Setiger 34. 
16. Notoseta from setiger one. 17. Neuropodial hook from posterior set- 
iger. 18. Neuropodial hook showing boundaries of internal hood. 19. 
Hook with primary hood removed, dorsal view. 20. Everted proboscis: a. 



Two new spionids 391 

Material examined: Type material of the following: Prionospio 
alata USNM 17369; Prionospio plumosa USNM 19596; P. tread- 
welli, Allan Hancock Foundation; P. ornata Holotype: USNM 32698 
Paratype: USNM 32697. Other material includes collections from the 
following: PERU coast; Mayaguez, PUERTO RICO; FLORIDA-Cedar 
Key, Clearwater Beach ( 4 sta. ) , Alligator Harbor ( 8 sta. ) , off shore Pan- 
ama City (4 sta.). North Bay near Lynn Haven (4 sta.), Gasparilla Is- 
land, Horseshoe Point; Grand Isle, LOUISIANA (76 samples); Port Ar- 
ansas, TEXAS; Beaufori:, NORTH CAROLINA; Yorktown, VIRGINIA; 
MALDIVES; Gold Coast of W. AFRICA; Capetown, S. AFRICA; Con- 
cepcion Bay, CHILE; PORTUGUESE GUINEA. 

Diagnosis: Body up to 94 mm in length. Prostomium spindle-shaped, 
inclosed by lateral wings formed by fusion of peristomium and first seg- 
ment. Three pairs of pinnate branchiae beginning on setiger one. Neuro- 
podial multidentate, hooded hooks begining on setiger nine; notopodial 
hooks beginning posterior to setiger 19. Pygidum with three anal cirri; 
two shorter laterals and a longer, unpaired dorsal cirrus. 

Description: The prostomium varies slightly in shape, from a very nar- 
row, tapered cylinder to a spindle-shaped structure, widest in the area of 
the area of the eyes. In the adult, there may be zero to four well-defined 
prostomial eyes and occasionally there are two larger area of diffuse pig- 
ment. The peristomium is fused dorsally and laterally with setiger one, 
forming an envelope inclosing the prostomium. There are two large wing- 
like dorsal peristomial expansions which may be closely applied to the 
prostomium or extended laterally (Fig. 12). 

The parapodia of segment one are absent. The palps are rarely pres- 
ent, being extremely deciduous; they are ventrally grooved and often 
possess a conspicuous basal sheath. The three pair of branchiae are pin- 
nate and very often missing. In the majority of cases where gills are 
present, one or more are in some stage of regeneration resulting in ex- 
treme variation in both number of pinnae and proportion of the shaft 
which is pinnate. In some instances the regenerating gill is completely 
smooth. The gills are variable in length and may reach from setiger two 
to setiger 15. The branchiae begin on the first setiger which is homolo- 
gous to setiger two in species of Prionospio. In some instances, a thread- 
like filament may arise at the base of the third pair of gills. There is a 
conspicuous dorsal ridge connecting the two gill bases on setiger one 
(Fig. 12). 

The anteriormost parapodial lamellae are well-developed and notopo- 
dial lamellae of setigers one to five are lanceolate (Fig. 13). The cor- 



dorsal view b. ventral view c. anterior view (not drawn to scale). 21. 
Pygidium, pigmentation shown by stipling. Figs. 16-19, scale A; Figs. 14, 
21, scale B; Fig. 15, scale C. 



392 Proceedings of the Biological Society of Washington 

responding neuropodial lamellae are somewhat smaller and less foliaceous. 
Posterior to setiger five, dorsal and ventral lamellae become increasingly 
rounded until they are somewhat similar in shape, though those of the 
neuropodium remains lower (Fig. 14). Progressively the notopodial 
lamellae become thin and long until posteriorly they are increasingly 
acuminate (Fig. 15). 

Anterior setae are all capillary and similar in appearance. Notopodial 
setae appear to be in three rows, the posterior row containing the long- 
est setae. The sheath often gives the setae a bilimbate appearance (Fig. 
16). Granulations are apparent. In this species, however, they often ap- 
pear on the sheath rather than the shaft which is where they are usually 
found. At setiger nine, neuropodial setae change abruptly with the ap- 
pearance of multidentate hooded hooks accompanied by long, thin com- 
panion setae (Fig. 17). There are one to three sabre-setae at the ventral 
edge of the setal fascicle. Notopodial hooks appear posterior to setiger 
19. Both neuro- and notopodial hooks have a large, clear, primary hood 
and a smaller, internal, secondary hood. The latter is closely applied to 
the setal shaft and is heavily striated (Figs. 17, 18, 19). 

On the dorsal surface between parapodia of a number of specimens, 
beginning about setiger 20, there are large, clear hyalinelike circles lo- 
cated dorsally with small dots in the center. These circles are accom- 
panied by pouches or thin membranes. The function of these structures is 
still unknown. 

The proboscis is bilobed and cylindrical (Fig. 20). The pygidium 
bears three anal cirri, a short lateral pair and a single, mid-dorsal, long, 
thin one (Fig. 21). 

Biology: This species has been dredged from substrata composed of 
mud, mud and clay, and mud and sand. It has been found in thin-walled 
tubes of mud ( Berkeley and Berkeley, 1952 ) and tubes of fine mud and 
clay ( Wesenberg-Lund, 1949). The only tubes observed on Caribbean 
specimens are transparent, fibrous and covered by relatively large quartz 
grains. P. pinnata is eurybathyal; found at depths ranging from less than 
three to 1300 m. 

Genus Aonides Claparede, 1864 

Aonides Claparede, 1864. Type species by monotypy: A. auricularis 
Claparede, 1864, [A. oxycephala (Sars, 1862)1. 

Paranerine Czemizvsky, 1881. Type species by monotypy: Nerine oxy- 
cephala Sars, 1862. 
Etymology: Gender: feminine; Aonides — L. — the Boeotian Women, 

i.e., the Muses. 

Diagnosis: Prostomium conical, tapered both anteriorly and posteriorly. 
Peristomium not well-developed. Branchiae cirriform, separate from dor- 
sal lamellae, beginning on setiger two and present on a variable number 
of anterior setigers. Hooded hooks present in both notopodia and 
neuropodia, bidentate or tridentate. Pygidium with anal cirri. Pelagic 



Two new spionids 393 

and lecithothrophic development. Spherical egg with thick membrane 
and short-headed sperm with tapered acrosome (Hannerz, 1956). 

Aonides mayaguezensis new species 

(Figs. 22-33) 

Diagnosis: Aonides with 15 to 16 pairs of branchiae. Body up to 10 
mm in length. Bidentate hooded hooks beginning in neuropodium on 
setigers 19 to 23 and in notopodium on setigers 21 to 24. With four py- 
gidial cirri, a shorter ventral pair and a longer dorsal pair. 

Material examined: Three specimens collected at a depth of 10 feet 
off Mayaquez, Puerto Rico, August, 1963. Holotype: USNM 39485; 
Paratype: USNM 39486. 

Description: The prostomium is elongate, thin and slightly wider in 
the region of the eyes (Fig. 22). There is an elevation in the posterior 
half, at the end of which is a digitiform process or occipital tentacle, 
which extends to or slightly beyond setiger one (Fig. 23). The pro- 
stomium terminates at the level of the first setiger. There are four eyes, 
the posterior pair being closer together than those of the anterior pair. 
The posterior pair is located on the prostomial elevation and is easily 
seen. The anterior pair, however, is found near the base of the elevation 
and is difficult to detect dorsally ( Figs. 22 and 23 ). The peristomium sur- 
rounds the posterior one-third to one-half of the prostomium giving a 
slight hoodlike appearance. 

The anterior branchial region is slightly flattened dorsoventrally. There 
are 15-16 pairs of branchiae, beginning on setiger two. The gills are 
almost as long as the body width generally longer than the dorsal lamellae, 
and are held erect dorsally. The first pair is only slightly, if at all, shorter 
than those following. The last pair, however, is less than half as long as 
the preceding and somewhat thinner. 

The parapodial lamellae of setiger one differ considerably from those of 
the following parapodia ( Fig. 24 ) . The dorsal lamella is broad, low and 
subtriangular. The neuropodial lamella, however, is much narrower, al- 
most digitiform. In the following paraodia the notopodial lamellae become 
increasingly foliaceous and the neuropodial lamellae increase slightly in 
width ( Fig. 25 ) . At about the level of setiger 10, the notopodial lamellae 
begin decreasing in size ( Fig. 26 ) until they are nearly the same height as 
those of the neuropodia, though they remain somewhat broader through- 
out the body length ( Fig. 27 ) . The decrease in size is quite abrupt in the 
postbranchial region. 

Anterior setae are capillaries and are arranged in two rows. In the 
neuropodia the anterior series contains shorter, thicker setae with a 
slightly wider sheath than those of the posterior row (Figs. 28, 29). There 
is an obvious sheath but the presence of one or two "limbations" depends 
on the orientation of the seta (Fig. 30). Granulations are evident on the 
majority of capillaries. In a notopodial fascicle there are two to three dor- 
sal, extremely thin, non-sheathed capillaries but the others are similar in 



394 Proceedings of the Biological Society of Washington 




Figs. 22-26. Aonides mayaguezensis new species. 22. Dorsal view, an- 
terior end. 23. Lateral view, anterior end. 24. Setiger one. 25. Setiger six. 
26. Setiger 10. Figs. 22, 23, 25, 26, scale B; Fig. 24, scale C. 



Tivo new spionids 395 

appearance and arrangement to those of the neuropodia. Bidentate, 
hooded neuropodial hooks appear on setiger 19-23 (19, on type-speci- 
men) (Fig. 31). Notopodial hooks of the same type appear on setigers 
21-24 (21, on type-specimen) (Fig. 32). There is no obvious secondary 
hood. The two teeth are widely separated and there are faint striations on 
the primary hood. 

The pygidium is drawn into four anal cirri (Fig. 33). Those of the 
ventral-most pair are short but about the same thickness as the longer 
dorsal pair. 

Discussion: Two species have been added to Aonides since the revision 
of the genus by Pettibone (1963). Of the species possessing bidentate 
hooded hooks, A. mayaguezensis shows a close affinity to A. californiensis 
Rioja, having similar branchiae, but they differ in the first appearance of 
neuro- and notopodial hooded hooks. Also, in A. californiensis the lamellae 
of setiger one are low and rounded whereas in the new species the ventral 
lamellae of this setiger are tapered, almost digitiform. Type material for 
A. californiensis no longer exists ( Dra. Maria Elena Casa Murioz, personal 
correspondence) leaving the original description as the only source of 
comparative information. On this basis, A. mayaguezensis is considered 
a separate species. 

KEY TO THE SPECIES OF AONIDES 

1. Hooded hooks bidentate 2 

1. Hooded hooks tridentate 



A. paucibranchiata Southern, 1914, Ireland. 

Second tooth of hooded hooks with longitudinal grooves, giving a 

tripartite appearance. Pygidium six anal cirri 

A. trifidus Estcourt, 1967, New Zealand ( Estuarine ) . 



2. Second tooth of hooded hooks lacking longitudinal grooves. Pygid- 

ium with four anal cirri 3 

3. Branchiae 20-30 pairs 

A. oxycephala (Sars, 1862), Mediterranean, Norway. 

3. Branchiae fewer than 20 pairs 4 

4. Neuropodial hooks beginning on setigers 19-23; notopodial hooks 

beginning on setigers 21-24. Branchiae 15-16 pairs 

A. mayaguezensis new species, Puerto Rico. 

4. Neuropodial hooks beginning on setiger 40; notopodial hooks begin- 

ing on setiger 35. Branchiae 13-14 pairs 

A. californiensis Rioja, 1947, Baja, California. 



In this paper, Aonides notoseta Storch, 1966, from the Red Sea, is not 
treated as a species of Aonides as it does not seem to conform to the ge- 
neric diagnosis. According to the original description, the principal charac- 
ters it shares with other species of Aonides are presence of anterior bran- 
chiae separate from the notopodial lamellae, hooks absent in anterior 
setigers and presence of anal cirri; none of these being diagnostic generic 
characters. Other characters which suggest that it does not belong in this 



396 Proceedings of the Biological Society of Washington 




Figs. 27-33. Aonides mayaguezensis new species. 27. Setiger 26. 28. 
Short neuroseta from setiger two. 29. Long neuroseta from setiger two. 30. 
Three neurosetae from setiger three. 31. Neuropodial hooded hook from 
setiger 26. 32. Notopodial hooded hook from setiger 26. 33. Pygidium. 
Figs. 28-32, scale A; Figs. 27, 33, scale C. 



Two new spionids 397 

genus are branchiae beginning on setiger three, hooks with a node or 
swelling on the shaft ( previously unknown among spionids ) , threadlike — 
"fadig" — ^lamellae and prostomial shape as figured in the original de- 
scription. No further designation exclusion from Aonides will be attempted 
until type material has been examined. 

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Spionidae to Sabellariidae. Proc. Acad. Nat. Sci. Phila., 75: 
179-259. 

Okuda, S. 1936. Spioniform polychaetes from Japan. J. Fac. Sci. Hok- 
kaido Univ. Zool., ser. 6, 5: 217:254. 

. 1937. Annehda Polychaeta in Onagawa Bay and its vicinity. 

Polychaeta Sedentaria. Sci. Rep. Tohoku Imp. Univ., ser. 4. 
Biol, 12(1): 45-69. 

Pettibone, M. H. 1963. Revision of some genera of polychaete worms of 
the family Spionidae, including the description of a new spe- 
cies of Scolelepis. Proc. Biol. Soc. Wash., 76: 89^104. 

Reish, D. 1959. An ecological study of pollution in L. A., Long Beach 
Harbors, Calif. Allan Hancock Fdn. Publ, Occ. Pap., 22: 1- 
119. 

. 1961. A study of benthic fauna in a recently constructed boat 

harbor in S. Calif. Ecology, 42( 1 ) : 84-91. 

RiojA, E. 1947. Estudios anelidologicos XVII. Contribucion al conocim- 
iento de los anelidos poliquetos de Baja California y Mar de 
Cortes. Anal. Inst. Biol., Mex., 18( 1 ) : 197-224. 

Sars, M. 1861. On armelids laegten Nerine og dans norske Arter. Vid- 
ensk. Selsk. Christiania, Forh., pp. 59-67. 

Shepherd, W. M. 1964. A guide to some Tomales Bay polychaetes. Univ. 
of the Pac. Pac. Marine Sta. NONR-3002 ( 02 ) Project NR 
104-626. viiiandpp. 1-34 (Mimeo.). 

SoDERSTROM, A. 1920. Studicn iiber die Polychaetenfamilie Spionidae. 
Inaug. Diss. Uppsala, 286 pp. 

Southern, R. 1914. Archiannelida and Polychaeta. In Clare Island Sur- 
vey. Pt. 47. Proc. Roy. Irish Acad. Dublin, 31: 1-160. 

Storch, Volker von. 1966. Drei neue Polychaeten aus dem Litoral des 
Roten Meeres. Kieler Meeres., 22(2): 171-175. 

Tebble, N. 1955. The polychaete fauna of the Gold Coast. Bull. Brit. 
Mus. (Nat. Hist.) Zool., 3(2): 61-148. 

Treadwell, a. 1931. Three new species of polychaetous annelids from 
Chesapeake Bay. Proc. U. S. Nat. Mus., 79, No. 2867: 1-5. 

Weese, a. O. 1933. The annelids of a marine sere. Acad. Sci. Oklahoma, 
13(3): 18-21. 

Wesenberg-Lund, E. 1949. Polychaetes of the Iranian Gulf. Danish Sci. 
Invest. Iran, Copenhagen, 4: 247-400. 

Wu, B. L. AND H. Chen. 1964. A new species of polychaete worm of the 
family Spionidae from Sisha Islands, with a review of the genus 
Prionaspio Mahngren, 1867. Acta Zool. Sinica., 16(1): 54- 
60. 



400 Proceedings of the Biological Society of Washington 



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402 3 October 1969 



PROCEEDINGS /^^^^^''^^ 

OF THE / 

BIOLOGICAL SOCIETY OF WASHINGTON ^ ^ 



^kmN^\^ 



A NEW JACKAL {CAmS ADUSTUS) FROM THE SUDAN 

By Henry W. Setzer 
Smithsonian Institution, Washington, D.C. 

In 1961 a detachment from the U.S. Naval Medical Research 
Unit No. 3 (NAMRU3) stationed in Cairo, Egypt, was detailed 
to the Sudan to conduct epidemiological studies in the vicinity 
of Malakal. During these field studies a series of jackals ( Canis 
adustus) was obtained which have proven to be distinct at the 
subspecies level from other populations of jackals in East 
Africa. 

All measurements given are in millimeters and capitalized 
color terms are from Ridgway, "Color Standards and Color 
Nomenclature," 1912. 

To Dr. Harry Hoogstraal and his associates in NAMRU3 and 
to the officials of the Field Museum of Natural History, Chi- 
cago, Illinois, I would like to express my thanks for the opportu- 
nity of describing: 

Canis adustus namrui new subspecies 

Holotype: Adult male, skin and skull. Field Museum of Natural His- 
tory, number 93856 from 1 mile south of Tir, Paloich District, Upper Nile 
Province, Republic of the Sudan. Obtained 3 March 1961 by Harry 
Hoogstraal, original number 13496. 

Specimens examined: Thirteen, from: 1 mi. W Tir, 2(1 ^ , 1 9 ); 1 mi. 
S Tir 5 ( 1 $,4 2 2 ) ; 12 mi. W Paloich, 1 male; Taufikia, Malakal, 1 $ ; 
10 mi. S. Paloich, 1 $ ; 5 mi. S Niayok, 1 $ ; 5 mi. S. Paloich, 1 $ ; 12 mi. 
N Malakal, 1 2 . 

Diagnosis: Reddish coloration of muzzle absent; general overall color 
grayish tan with strong admixture of black on back; belly color near Pink- 
ish Cinnamon; anterior parts of forelegs near Buckthorn Brown and lack- 
ing any pronounced anterior stripe; cheeks, post- and subauricular areas 
white; backs of ears same color as back. Tail short, black dorsally and 
terminally all around. Skull small for the species; teeth relatively small; 
interpterygoid space lyre-shaped nasals relatively broad posteriorly. 

29— Proc. Biol. Soc. Wash, Vol. 82, 1969 (401) 



OCT 3 1969 



402 Proceedings of the Biological Society of Washington 

Comparisons: Compared with Canis adustus kaffensis (Neumann, 
, 1902), as known by specimens from Agarasalam, Sidamo District, Ethio- 
pia, C. a. namrui differs in generally paler color; gray, rather than reddish 
brown, muzzle; absence of dark streak on anterior face of forearm and 
markedly paler legs and under parts. The skull is smaller in all respects; 
the interpterygoid space is lyrate rather than straight sided; and the zygo- 
rnatic arches are less flaring laterally. 

From Canis adustus bweha Heller, 1914, as known from Guasu Ngishu 
Plateau, Kenya, C. a. namrui differs in markedly paler color and ap- 
parently smaller size. The auditory bullae are larger and more inflated; 
the teeth are smaller; the zygomatic arches are less flaring laterally; and 
markedly less inflation occurs in the frontal area of the skull. 

C. a. namrui differs from Canis adustus notatus Heller, 1914, as known 
from Masindi, Uganda, in markedly paler color and in somewhat smaller 
size. The skull is markedly smaller in every respect, especially in the size 
and degree of inflation of the auditory bullae, the size of the cheek teeth 
and in the length of the bony palate. 

Measurements: Holotype. — Total length 925; length of tail 276; length 
of hind foot 151; length of ear ?; greatest length of skull 151.5; condyloin- 
cisive length 146.2; length of audital portion of auditory bulla 22.5; great- 
est breadth across zygomatic arches 75.9; least postorbital breadth 25.2; 
least interorbital breadth 25.9; greatest length of nasals 55.5. 

Type series: Measurements with means and extremes of 4 males and 7 
females, are respectively: Total length 1002.5 (925-1060), 946.3 (904- 
1002); length of tail 286.8 (276-310), 289.1 (278-316); length of hind 
foot 160.5 (151-172), 154.7 (147-168); length of ear from notch 73.0 
(67-76), 72.7 (68-79); greatest length of skull 150.9 (142.2^-156.0), 
140.8 (137.2^144.3); condyloincisive length 146.5 (140.1-150.3), 136.3 
(131.6-144.8); length of audital portion of auditory bulla 23.1 (22.2- 
23.8), 21.4 (20.3-22.9); greatest breadth across zygomatic arches 
76.6 (75.8-78.2); 69.7 (66.2^74.0); least postorbital breadth 27.3 (24.7- 
31.3), 27.2 (23.5-31.0); least interorbital breadth 25.4 (23.1-27.3), 22.1 
(20.3-23.2); greatest length of nasals 57.3 (53.4-61.6), 53.9 (45.4-61.4). 

Remxirks: No intergradation can be demonstrated between C. a. namrui 
and any of the adjacent subspecies, but this is undoubtedly owing to lack 
of specimens from intermediate areas rather than its total absence. Inter- 
grades, when found, should be readily distinguishable if coloration is at 
all important. 

Color in this new subspecies is so markedly different from other jackals 
from surrounding areas that specimens are readily identifiable on this 
character alone. 

Comparisons with subspecies of Canis adustus other than those above 
are not necessary inasmuch as these three subspecies geographically stand 
between C. a. namrui and other named subspecies. 

Etymology : Canis adustus namrui is named in recognition of the exten- 
sive epidemiological work done in northeastern Africa by the staff of Naval 
Medical Research Unit No. 3. 



^ 13 

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Vol.1 HH 3 October 1969 



PROCEEDINGS 
OF THE 

BlULuv^.v-AL SOCIETY OF WASHINGT 



BATHYCONCHOECIA DEEVEYAE, A HIGHLY 

ORNAMENTED NEW SPECIES OF OSTRACODA 

(HALOCYPRIDIDAE) FROM THE PERU-CHILE 

TRENCH SYSTEM 

By Louis S. Kornicker 
Smithsonian Institution, Washington, D.C. 

Deep-sea benthonic organisms were collected from the Peru- 
Chile Trench System on Cruise 11 of the R/V Anton Bruun, 
which was sponsored by the National Science Foundation. 
Among the ostracods was a single specimen of a most unusual 
halocyprid ornamented with 7 long spines. This new species is 
described below. 

SUBORDER HALOCYPRIDINA 

HALOCYPRIDIDAE 

Bathyconchoecia Deevey, 1968 

Type-species: Bathyconchoecia paulula Deevey, 1968. 

Appendages of the new species described herein differ from those de- 
scribed in the diagnosis of the genus by Deevey ( 1968 ) in several charac- 
ters. The most important of these is the presence of only 6 claws on the 
furca instead of the usual 8. Because the presence of only 6 furcal claws is 
probably a juvenile character, the present species is included in Bathycon- 
choecia. 

Bathyconchoecia deeveyae new species 
Plate 1, Figures 1, 2 

Holotype: USNM 123335; carapace preserved in alcohol; appendages 
on slide in glycerine; specimen unique. 

Type-locality: Pacific Ocean, 07°53'S, 80°30'W; Bottom depth 520' m; 
Station 85, Cruise 11, R/V Anton Bruun, 14 October 1965; collected in 
benthic trawl, 520 mesh net, by Dr. R. J. Menzies. 

Etymology: The species is named for Dr. Georgiana B. Deevey. 

Description of juvenile: Shell surface with minute pits which are es- 
pecially apparent under phase illumination (fig. la); incisure deep with 
rounded inner margin; rostrum with pointed tip extending in front of and 

30— Prog. Biol. Soc. Wash., Vol. 82, 1969 (403) 




404 Proceedings of the Biological Society of Washington 




A new ostracod 405 

below incisure, and bearing long hollow spine; long hollow spine also pres- 
ent at dorsal margin on posterior of right valve; each valve with short 
spine near dorsal margin anterior to valve middle, and short spine near 
valve middle below its center; asymmetrical glands present symmetrically 
near posterodorsal corner of each valve. 

Size: Length without spines — 1.12 mm; length including spines — 1.61 
mm; height without spine 0.89 mm. 

First antenna (fig. Id-f): Shaft short with segments indistinct on ex- 
amined specimen. Brushlike structure on 3rd segment with about 148 
filaments in about 8 rows, each with 18 filaments. Long stout spinous 
bristle present distal to brushlike structure. End segment with 1 long and 
2 shorter bristles or filaments, latter bristles about same length as spinous 
bristle of adjacent segment. 

Second antenna (fig. Ig-j); Endopodite; 1st segment with 2 distodorsal 
bristles; end segment or segments with a total of 5 setae and filaments. 
Exopodite: distal margins of 3rd and 4th segments with comb of fine 
teeth; bristles of segments 2-8 with sparse natatory hairs; 9th segment 
with 3 bristles, 1 long, 1 medium, 1 short. 

Mandible (fig. 2a-e): Coxa pars incisiva with 5-6 teeth present on 
fingerlike posterior process; a single small tooth present anterior and prox- 
imal to large anterior tooth of pars incisiva; proximal tooth list furnished 
with 4-5 saw-like teeth; distal tooth list furnished with 8-9 teeth; anterior 
corner of coxa with spinous short, stout bristle; numerous long spines pres- 
ent between bristle and proximal tooth list. Basale: distal edge of endite 
with 6 large, triangular teeth followed by 2 recurved processes; anterior 
edge of endite with 1 long bristle; lateral surface of endite with 3 long 
bristles; single medial bristle present on segment; 2 transparent bristles 
with marginal hairs representing exopodite ( ? ) present near dorsal margin 
of segment. Endopodite: 1st segment with single distodorsal bristle and 2 
ventral bristles near middle; 2nd segment with 3 distodorsal bristles and 1 
ventral bristle; end segment with 7 bristles, 4 short, 2 medium, and 1 ex- 
tremely long; fine hairs present on medial surface of end segment. 

Maxilla ( fig. Ik-m ) : Precoxale endite with 6 bristles; coxale endite 
with 12 bristles, 6 on each lobe. Endopodite: anterior margin of 1st seg- 
ment with 1 bristle near middle and 4 distally; posterior margin with 3-^ 
bristles proximally and 1 distally; end segment with 2 short stout curved 
claws and 4 short slender bristles between the claws. 

Fifth limb (fig. 2f ) : Exopodite: 1st segment with extremely long bristle 
on dorsal margin reaching past bristles on end segment of exopodite; 2nd 
segment with 2 bristles near middle of ventral margin and 1 on dorsal 
margin; end segment with 2 clawlike bristles and 1 slender bristle. Endo- 



Plate 1. — Bathyconchoecia deeveyae Kornicker, carapace, total length 
1.61 mm: a, dorsal view; b, right lateral view; c, ventral view. (Illustra- 
tion prepared by Carolyn Bartlett Cast. ) 



SWiVH-n^:'? ijQjS 1989 



406 Proceedings of the Biological Society of Washington 




Figure 1. — Bathyconchoecia deeveyae Kornicker, USNM 123335: a, 
detail of surface pits on carapace in antero-dorsal area near hingement; b, 
sketch showing position of appendages, anterior to right; c, dorsal view of 
"c", anterior to right. First antenna: d, complete limb; e, f, distal ends. 
Second antenna: g, left limb, medial view; h, distal part left endopodite, 
medial view; i, right limb, lateral view; j, distal part right endopodite, 
lateral view. Maxilla: k, complete limb; precoxale endite; m, coxale en- 
dites, anterior to right. (Same magnification, in microns: a; b, c; d, g, i, k; 
e, f ; h, j, 1, m. ) 



A new ostracod 



407 




Figure 2. — Bathyconchoecia deeveyae Kornicker, USNM 123335, man- 
dible: a, complete right limb, lateral view; b, detail of basale and coxale 
on "a"; c, left mandible, medial view; d, detail of basale of "c", medial 
view. Posterior limbs: f, 5th limb; g, 6th limb; h, 7th limb; i, right lamella 
and middle bristle of furca. (Same magnification, in microns: a, c, f, g, i; 
h;b,d,e.) 



podite with 2 claw-like bristles. Epipodial appendage with 12 bristles 
in 3 groups of 4 bristles separated by a space. 

Sixth limb (fig. 2g) : Exopodite: 1st joint with 3 short bristles and 1 ex- 
tremely long distodorsal bristle reaching just past bristles of 4th joint of 



408 Proceedings of the Biological Society of Washington 

exopodite; 2nd joint with 2 distoventral bristles; 3rd joint with 1 ventral 
and 1 dorsal bristle; 4th joint with 3 long bristles, ventral bristle more 
slender than others. 

Seventh limb (fig. 2h) : End segment with 2 long subequal bristles. 

Furca (fig. 2i) : Each lamella with 6 claws; claw 1 longer than claw 2; 
claw 2 only slightly longer than claws 3-6; a long spinous bristle present 
between lamellae following last claw; surface of lamellae with short spines. 

Pigmentation: Areas with black pigmentation present in preserved 
specimen posterior to 1st segment of each 1st antenna and anterior to 
heart ( pi. la, fig. Ic ) . 

Rod-shaped organ: Not observed with certainty. 

Ecology: The specimen was collected on the bottom at a depth of 520 
m. Ahydrocast (No. 84; 07°50' S, 80°29' W; 13 October 1965) obtained 
by the R/V Anton Bruun in the vicinity of Station 85 where the ostracod 
was collected gave the following water properties at depth of 500 m: 
Temp.— 7.74°C; Sal.— 34.637 parts per thousand; 0^0.15 ml./l (Men- 
zies and Chin, 1966). 

Comparisons: The spinosity of the carapace separates this species from 
other members of the genus. 

Literature Cited 

Deevey, G. B. 1968. Bathyconchoecia, a new genus of pelagic ostracod 
(Myodocopa Halocyprididae) with six new species from the 
deeper waters of the Gulf of Mexico. Proc. Biol. Soc. Wash., 
vol. 81, pp. 539-570, 13 figs. 

Menzies, R. J. AND Chin, E. 1966. Cruise Report, Research Vessel Anton 
Bruun, Cruise 11. Special report number 1, Marine Labora- 
tory, Texas A&M University, Galveston, Texas, 16 pp., 15 figs., 
2 tables. Append. I, II, III. [Statement in report to effect that 
it does not constitute a publication.] 



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